Bulletin ILLINOIS NATURAL HISTORY SURVEY JtSUllQTtn Printed by Authority of the State of Illinois The Amphibians and Reptiles of Illinois PHILIP W. SMITH STATE OF ILLINOIS Otto Kerner, Governor DEPARTMENT OF REGISTRATION AND EDUCATION William Sylvester White, Director NATURAL HISTORY SURVEY DIVISION Harlow B. Mills, Chief 0£C 2 1 MM !' SURVEi! ,^w^- ILLINOIS NATURAL HISTORY SURVEY Bulletin ^^ Volume 28, Article 1 ^^ Printed by Authority of November, 1961 the state of Illinois The Amphibians and Reptiles of Illinois PHILIP W. SMITH STATE OF ILLINOIS Otto Kerner, Governor DEPARTMENT OF REGISTRATION AND EDUCATION William Sylvester White, Director NATURAL HISTORY SURVEY DIVISION Harlow B. Mills, Chief Urbana, Illinois STATE OF ILLINOIS Otto Kerner, Governor DEPARTMENT OF REGISTRATION AND EDUCATION Wii-LiAM Sylvester White, Directoi BOARD OF NATURAL RESOURCES AND CONSERVATION William Sylvester White, Chairman; A. E. Emerson, Ph.D., Biology; Walter H. Newhouse, Ph.D., Geology, Roger Adams, Ph.D., D.Sc, Chemistry; Robert H. Anuerson, B.S.C.E., Engineering; W. L, Everett, E.E., Ph.D. Representing the President of the University of Illinois; Delyte W. Morris, Ph.D., President oj Southern lllinoi. University NATURAL HISTORY SURVEY DIVISION, Urbana, Illinois SCIENTIFIC AND TECHNICAL STAFF Harlow B. Mills, Ph.D.. Chiel Bessie B. East, M.S., Assistant to the Chu! Section of Economic Entomology Georoe C. Decker, Ph.D., Principal Scientist and Head J. H. BiGOER M.S., Entomologist L. L. English, Ph.D., Entomologist W. H. LucKMANN, Ph.D., Entomologist Willie N. Bruce, Ph.D., Entomologist John P. Kramer, Ph.D., Associate Entomologist Richaru J. DvsART, Ph.D., Associate Entomologist RoNAiD H. Meyer, M.S., Assistant Entomologist Reginald Roberts. M.S., Technical Assistant James W. Sanford, B.S., Technical Assistant Earl Stadelbacher, B.S., Technical Assistant William C. Move, M.S., Technical Assistant Sue E. Watkins. Technical Assistant H. B. Petty. Ph.D., Extension Specialist in Entomology* Stevenson Moore, III, Ph.D., Extension Specialist in Entomology* Zen as B. Noon. Jr., M.S., Research Assistant* Clarence E. White, B.S., Instructor in Entomology Extension* CosTAS Kouskolekas, M.S., Research Assistant* A.MAL Chandra Baxerjee, M.S., Research Assistant* Victor T. Williams, B.S., Research Assistant* Section of Faunistic Surveys and Insect Identification H. H. Ross, Ph.D., Principal Scientist and Head Milton W. Sanderson, Ph.D., Taxonomist Lewis J. Stannard, Jr., Ph.D., Taxonomist Philip W. Smith. Ph.D., Associate Taxonomist Leonora K. Glovd, M.S., Assistant Taxonomist H. B. Cunningham, M.S., Assistant Taxonomist Ruth P. Cash, Technical Assistant Wayne Becker, B.S., Research Assistant John M. Kincsolver. Ph.D., Research Associate* John D. Unzicker. Research Assistant* Talaat K. Mitri, M.S., Research Assistant* Section of Aquatic Biology George W. Bennett. Ph.D., Aquatic Biologist and Head William C. Starrett. Ph.D., Aquatic Biologist R. W. Larimore. Ph.D., Aquatic Biologist David H. Buck, Ph.D., Associate Aquatic Biologist Robert C. Hiltibran. Ph.D.. Associate Biochemist Donald F. Hansen, Ph.D., Associate Aquatic Biologist William F. Childers. M.S., Assistant Aquatic Biologist Marifran Martin, Technical Assistant Robert D. Crompton, Field Assistant Michael J. Duever, Field Assistant Rollin D. Andrews, III, B.S., Field Assistant* Leslie L. Layton, Field Assistant* Section of Aquatic Biology—continued Charles F. Tiioits, III, A.B., Research Associate* Section of Applied Botany and Plant Pathology J. Cedric Carter, Ph.D., Plant Pathologist and Head J. L. Forsberg, Ph.D., Plant Pathologist G. H. BoEWE, M.S., Associate Plant Pathologist Robert A. Evers, Ph.D., Associate Botanist Robert Dan Neely', Ph.D., Associate Plant Pathologist E. B. Himelick, Ph.D., Associate Plant Pathologist Waltlr Hartstirn, Ph.D., Assistant Plant Pathologist D. F. ScHOENEWEiss. Ph.D., Assistant Plant Pathologist Anne Rohinso.n, M.A., Technical Assistant Section of Wildlife Research Thomas G. Scott, Ph.D., Wildlife Specialist and Head Ralph E. Yeatter, Ph.D.. Wildlije Specialist F. C. Bellrose, B.S., Wildlife Specialist H. C. Hanson, Ph.D., Associate Wildlife Specialist Richard R. Graber, Ph.D., Associate Wildlife Specialist Ro.n'ald F. Labisky, M.S., Associate Wildlife Speci::list Gle.n C. Sanderson, M.A., Associate Wildlife Specialist Marjorie J. Schlatter, Technical Assistant D. G. Rose, B.S., Technical Assistant Howard Crum, Jr., Field Assistant Rexford D. Lord, D.Sc, Project Leader* Jack A. Ellis, M.S., Project Leader* Bobbie Joe Verts. M.S., Project Leader* Ralph J. Ellis, M.S., Project Leader* William L. Anderson, B.S., Assistant Project Leader* James A. Harper, M.S., Assistant Project Leader* David A. Casteel, B.S., Assistant Project Leader* Gerald G. Montgomery, M.S., Research Associate* P. J. Rao, B V.Sc, M.A., Research Assisant* Ann C. V. Holmes, B.S.. Research Assistant* T. U. Meyers, B.S., Research. Assistant* Stuart H. Mann, B.S., Research Assistant* Richard W. Lutz, M.W.M., Research Assistant* Richard D. Andrews, M.S., Field Mammalogist* Keith P. Dauphin, Assistant Laboratory Attendant* Section of Publications and Public Relations James S. .Ayars, B.S., Technical Editor and Head Blanche P. Voung, B.A., Assistant Technical Editor Edward C. Visnow, M.A., Assistant Technical Editor WiLMER D. Zeiir, Assistant Technical Photographer Technical Library Ruth R. Warrick, B.S., B.S.L.S., Technical Librarian Barbara S. McCrimmon, B.S., M.S.L.S., Assistant Tech- nical Librarian CONSULTANTS: Herpetology, Hobakt M. Smith, Ph.D., Professor of Zoology, University ol Hlinois; Parasitology, Norman D. Levine, Ph.D., Professor of Veterinary Parasitology and of Veterinary Research. University of Illinois: Wildlife Research. Wiliard D. Klimstra, Ph.D., Professor of Zoology and Director of Co-operative Wildlife Research. Southern Illinois University. Employed on co-operative projects with one of several aRenries: University of Illinois, Illinois Agricultural Extension Service, Illinois Dep.irtment of Conservation, National Science Foundation, United States Department of Agriculture, United States Fish and Wildlife Service, United States Public Health Service, and others. This paper is a contribution from the Section of Faunistic Surveys and Insect Identification. (08324—6M—10-60) <,^^feB 2 FOREWORD For the better part of a century the Bulle- tin has published research reports on the ecol- ogy, biology, and taxonomy of the flora and fauna of Illinois. The present number is a worthy continuance of these scientific contri- butions. It was prepared under those legal charges to the Board of Natural Resources and Conservation, the agency which controls, guides, and governs the activities of the Natu- ral History Survey, to "conduct a natural history survey of the State giving preference to subjects of educational and economic im- portance," and "To publish, from time to time, reports covering the entire field of zool- ogy and botany of the State." It is probable that no other groups of ani- mals have such staunch friends and adamant adversaries as do the amphibians and the reptiles, the subjects of this contribution. They are important to us in many ways, and, if they were not, "the characteristically human desire to investigate the unknown . . . provides an incentive to study them," as the author of this paper states. We humans are likely to be selfish in our dealings with the so-called lower animals and to measure the worth of these animals only in terms of their economic elect upon us. We find that many amphibians and reptiles add materially to the well-being of people—some by the destruction of harmful insects and rodents, others by affording additions to our diet, as with frog legs or (to the adventurous palate) rattlesnake flesh. Those people with an aesthetic bent find in amphibians and reptiles a great deal of grace and beauty. Those interested in curious things find their inquisitiveness satisfied among ani- mals which have no legs, or which can change color, or which breathe largely through their skins, or which go through generation after generation without really growing up. Those who look for antiquity find that giant amphib- ians and reptiles de^eloped, prospered, and passed on as living entities long before Nature began to experiment with the upright bipedal animals we call humans. Amphibians and reptiles are excellent organisms to study if one is interested in the variability of living things, as Dr. Philip W. Smith has amply demonstrated in this article. The movement of animals over the face of the earth in times past has intrigued biologists since the time of Aristotle. How did the pres- ent Illinois amphibians and reptiles reach the state? If they came from outside its limits, from what direction did they emigrate? What were the reasons for this movement? Illinois has proved to be a meeting place for many forms which apparently came from all directions, and many were unable to get past it or to invade it completely. Of the 94 species discussed here, only 25 are state-wide in their distribution. The other 69 have met limiting barriers somewhere in Illinois. For many species, parts of the state represent unin- habited islands, with the populations lapping at the shorelines on two, three, or four sides, but unable to overflow the highlands. For other species, parts of the state seem to repre- sent pools, small pockets of animals which cannot get out of circumscribed areas. The study on which the present report is based was initiated in June, 1947, and v;as one of the principal research projects of the Sec- tion of Faunistic Surveys and Insect Identifi- cation until June, 1953. The manuscript was completed in May, 1957, and was submitted for publication at that time. Budgetary reas- ons necessitated setting it aside until July, 1959. Although some revision has been neces- sary to bring the nomenclature and certain concepts up-to-date, the manuscript is approxi- mately the same as that submitted in May, 1957, and only the most pertinent of the recent literature has been incorporated. Hapi-Ovv B. Mills, Chief Illinois Natural History Survey Urbana, Illinois CONTENTS Acknowledgments ] Previous Herpetologicai Investigations 2 Illinois as a Herpetofaunal Region 4 Physical Features 4 Climatic Features 8 Biotic Provinces 8 Herpetofaunal Divisions 8 Presentation of Data 14 Systematic Account 15 Check List of the Amphibians and Reptiles of Illinois 15 Species Deleted From the Illinois List 18 Class Amphibia 21 Order Caudata (Salamanders) 22 Family Cryptobranchidae 25 Family Ambystomatidae 27 Family Salamandridae 39 Family Plethodontidae 41 Family Amphiumidae 61 Family Proteidae 61 Family Sirenidae 63 Order Salientia (Frogs and Toads) 66 Family Pelobatidae 69 Family Bufonidae 71 Family Hylidae 77 Family Ranidae 93 Family Microhylidae Ill Class Reptilia 113 Order Testudines (Turtles) 114 Family Chelydridae 117 Family Kinosternidae 122 Family Testudinidae 129 Family Trionychidae 154 Order Squamata 159 Suborder Sauria 1 59 Family Iguanidae 160 Family Anguidae 163 Family Teiidae 165 Family Scincidae 166 Suborder Serpentes 1 72 Family Colubridae 177 Family Crotalidae 263 Literature Cited 274 Index 287 Robert Kennicott 1835—1866 Harrison Carman 1858—1944 Karl P. Schmidt 1890—1957 Alvin R. Cahn 1892— Clifford H. Pope 1899— Fred R. Cagle 1915— Important contributors to the knowledge of the Illinois herpetofauna. The Amphibians and Reptiles of Illinois PHILIP W. SMITH The amphibians and reptiles comprise :wo classes of vertebrate animals. The class Amphibia includes three present-day or- ders, the Caudata (salamanders), Salientia (frogs and toads), and the Gymnophiona (caecilians). The class Reptilia includes four present-day orders, the Testudines (turtles), Squamata (lizards and snakes), iTrocodylia (crocodilians) , and Rhynchoce- phalia {Sphenodon). In the fauna of lUi- lois, amphibians are represented by sala- manders, toads, and frogs; reptiles by tur- des, lizards, and snakes. Altogether 109 dif- ferent kinds of these animals have been found in the state. In our natural economy the amphibians and reptiles, because of their predation on insects and destructive rodents, are on the side of man. Their exact role in the so- called balance of nature is at this time poorly known, but their existence is prob- ably of some importance to other organisms in the over-all fauna. To many people am- phibians and reptiles are fascinating and colorful creatures, and the characteristically human desire to investigate the unknown and to accumulate knowledge provides an incentive to study them. The increase in popularity in recent years of these animals is reflected by the number of people who now have pet turtles, lizards, and snakes. Four of our Illinois snakes are poisonous. Although these snakes are uncommon in most parts of the state, a few persons are bitten by them, and many persons worry needlessly because harmless species are mis- taken for poisonous species. It is advisable for everyone who spends much time out-of- doors to learn to distinguish the poisonous species ; this can be done readily with the information in this bulletin. A detailed study of the present-day Illi- nois amphibians and reptiles is bringing to light important data that help the scientist to look back into the past history of the earth and its life. Illinois has contributed especially significant data to biogeography, particularly in helping to disclose the move- ments of animals in and around Illinois dur- ing the advance and retreat of the glaciers of the Pleistocene. The objectives of this report on the am- phibians and reptiles of Illinois are three- fold. The first is to provide a critical re- view of the species and subspecies known to inhabit Illinois. The second is to present de- tailed distributional information for these animals in the hope that the data may con- tribute to the knowledge of the ecology and biogeography of Illinois. The third is to call attention to variation trends that I have discerned within Illinois and that will en- able future investigators to utilize character analyses for populations occurring in lim- ited parts of the state. ACKNOWLEDGMENTS The co-operation of several institutions and a great many persons has materially in- creased our knowledge of the herpetofauna of Illinois. The institutional and private collections that have made their holdings available to me are as follows: Chicago Academy of Sciences, Chicago, Illinois (CAS) ; Carthage College, Carthage, Illi- nois (CC) ; Chicago Natural History Mu- seum, Chicago, Illinois (CNHM); collec- tion of Charles W. Myers, St. Louis, Mis- souri (CM); Cornell University, Ithaca, New York (CU) ; Eastern Illinois Univer- sity, Charleston, Illinois (EIU); collection of Fred A. Shannon, Wickenburg, Arizona (FAS) ; field zoology collection accumu- lated at the University of Illinois by the late H. J. Van Cleave, Urbana, Illinois (HJVC) ; Illinois Natural History Survey, Urbana, Illinois (INHS); Illinois State Museum, Springfield, Illinois (ISM); col- lection of James C. List, Paducah, Ken- tucky (JCL) ; Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (MCZ); Nebraska State Museum, Lincoln, Nebraska (NSM) ; Prin- cipia College, Elsah, Illinois (PC) ; collec- tion of Richard A. Edgren, Chicago, Illinois (RAE); collection of Robert C. Schroder, Rock Island, Illinois (RCS) ; Rockford [1] Illinois Natural History Survey Bulletin Vol. 28, Art. 1 Museum of Natural History, Roclctord, Illinois (RMNH); collection of Sherman A. Minton, Indianapolis, Indiana (SAM); Southern Illinois University, Carbondale, Illinois (SIU); University of Illinois Mu- seum of Natural History, Urbana, Illinois (UIMNH); University of Michigan Mu- seum of Zoology, Ann Arbor, Michigan (UMMZ) ; University of Rochester, Roch- ester, New York (UR) ; United States Na- tional Museum, Washington, D. C. (USNM). I am indebted to the following persons for the privilege of examining material de- posited in the above mentioned collections: Esther Bennett, Fred R. Cagle, Doris M. Cochran, D. Dwight Davis, William E. Duellman, Richard A. Edgren, William E. Fahy, Howard K. Gloyd, Norman Hart- weg, Donald F. Hoffmeister, Robert F. Inger, Remington Kellogg, Alice Kibbe, James N. Layne, James C. List, Hymen Marx, Sherman A. Minton, Charles W. Myers, R. Earl Olson, Paul W. Parmalee, James A. Peters, Clifford H. Pope, Walter M. Scruggs, the late Karl P. Schmidt, C. Bertrand Schultz, Fred A. Shannon, Ho- bart M. Smith, Charles F. Walker, John F. Wanamaker, and T. Gilbert Wright. Sizable personal collections of specimens have been donated to the Illinois Natural History Survey by Jack Daugherty, Ed Keiser, Jr., Dave A. Langebartel, R. Earl Olson, Robert C. Schroder, Warren P. Sights, Jerry Staedeli, and Harlan D. Wal- ley. Specimens contributed from time to time by Richard W. Abbuhl, W. Leslie Burger, Donald J. Daleske, M. Max Hens- ley, James E. Huheey, Harold R. Hunger- ford, Allan H. Keith, R. Weldon Larimore, Ross J. Miller, Thomas E. Moore, William B. Robertson, Milton W. Sanderson, Robert Shoop, Lewis J. Stannard, and Kenneth L. Williams have been invaluable in filling in distributional gaps. Smaller lots of speci- mens and information have been contrib- uted by so many persons that they cannot be listed individually here. The assistance of each is sincerely appreciated. I wish to thank Howard K. Gloyd, who actually initiated this study, for relinquish- ing the results of his preliminary investiga- tions. I have profited by discussing with Hobart M. Smith, Herbert H. Ross, Sher- man A. Minton, and Roger Conant many of the problems encountered. I have been advised by several investigators on problems relating to their particular specialties, and acknowledgment for their assistance ap- pears in appropriate places throughout the text. For countless profitable consultations on editorial problems that arose during the writing of the manuscript and for reading proof, I owe a special gratitude to my asso- ciate, Mrs. Leonora K. Gloyd. For her accurate typing of the manuscript and for her vigilance in detecting inconsistencies, I am grateful to Mrs. Thelma H. Over- street. For illustrations, I am indebted to several persons, most of them formerly or presently on the staff of the Natural History Survey. The key illustrations were done by Joseph I. Leveque and Mrs. Alice Ann Prickett; the drafting by Miss Marguerite M. Ver- ley. Most of the photographs were taken by William E. Clark; smaller numbers were taken by Charles L. Scott, Robert E. Hesselschwerdt, and Harry Fisher. The photos by Mrs. Isabelle Hunt Conant (figs. 16, 89, 115, 149, 190, 210, and 229) are from the photographic collection of Roger and Isabelle Hunt Conant. The photograph of Robert Kennicott in the frontispiece is reproduced through the courtesy of the Chi- cago Academy of Sciences, that of Harrison Garman through the courtesy of Dr. Lee R. Townsend of Lexington, Kentucky, and that of Karl P. Schmidt through courtesy of the Chicago Natural History Museum. For editing the manuscript and for read- ing of the proof, I am grateful to James S. Ayars, Mrs. Diana R. Braverman, Mrs. Nira J. Nelson, and Mrs. Blanche P. Young. I am indebted to my former major pro- fessor, Hobart M. Smith, to my immediate superior, Herbert H. Ross, and to my wife, Dorothy M. Smith, for continuous help throughout the course of this study. PREVIOUS HERPETOLOGIGAL INVESTIGATIONS Three reports on the amphibians and rep- tiles of Illinois have appeared. One of these is a briefly annotated check list, one a key only, and one an excellent account, which is, however, more than 60 years old. Four pa- pers dealing with limited groups of the her- petofauna and approximately half a dozen papers treating the species of limited re- November, 1961 Smith: Amphibians and Reptiles of Illinois gions within Illinois have been published. The great majority of the references to Illi- nois in the literature are incidental, either citing specimens collected at localities in Illinois or merely stating that Illinois is within the range of the species. The earliest specific reference to an am- phibian or reptile in Illinois appears to be an observation by Thomas Say (1823) in the account of Long's expedition to the Rockies, just 5 years after Illinois became a state. Say records in his narrative that he observed a female of the then recently de- scribed Graptemys geographica laying eggs on the bank of the Ohio River near Shaw- neetown. Fifteen years later Holbrook (1838) described Emydoidea blandingi from the Fox River of northern Illinois. Geb- hard (1854) recorded the first amphibian for the state by citing a specimen of Hemi- dactylum [sic] from Illinois. Kennicott (1855) prepared the first regional list with his catalog of Cook County quadrupeds. His list records 12 amphibians and 21 rep- tiles still recognized as part of the Cook County fauna. Later Kennicott (1856) de- scribed Natrix kirtlandi from West North- field, and (1859) Virginia valeriae elegans from "southern Illinois," and Ophibolus ev- ansii from "central Illinois." During the next 25 years such authors as Cope, Cooper, Yar- row, Jordan, and Kennicott published pa- pers containing incidental references to the herpetofauna of Illinois. Davis & Rice (1883fl) published the first state list; their check list correctly recorded 29 species of amphibians and 54 reptiles, as well as a number of other species whose oc- currence is now known to be quite impossi- ble in Illinois. In the same decade Schneck (1880-1886), a physician and naturalist in Mount Carmel, was active in the Wabash River valley, and his several short notes in the American Naturalist recorded a num- ber of reptiles in southeastern Illinois. Cope (1888) described Lampropeltis triangulum syspila from Richland County and (1889) Hyla phaeocrypta from Mount Carmel. Harrison Garman (1889, 1890) recorded the animals occurring in the Mississippi River bottoms near Quincy and reported several species for the first time from the state. Hay (1881-1892), studying the her- petology of Indiana, contributed to the knowledge of the Illinois herpetofauna by citing Illinois localities for several species. Cope (1892) described Thamnophis sirtalis semifasciata from Des Plaines in northeast- ern Illinois. The synopsis by Harrison Garman (1892), until now the only comprehensive report on the Illinois amphibians and rep- tiles, credited correctly 31 species of am- phibians and 58 reptiles to Illinois, but un- fortunately perpetuated most of the errors of Davis & Rice. Hurter (1893-1911) added several species to the known fauna ot southwestern Illinois and, in his Herpetol- ogy of Missouri, described Chrysemys tre- leasei from Madison, St. Clair, and Monroe counties, Illinois. In the early part of the present century, Robert Ridgway was responsible for a great quantity of Illinois material which found its way into the United States National Mu- seum. Ridgway was also responsible for a number of erroneous records, presumably based on his recollections of species proba- bly misidentified in the field. Shelford (1913) cited a number of species for north- eastern Illinois in his Animal Communities of Temperate America. Gaige (1914) re- corded 16 species of amphibians and reptiles for Richland County, and Hankinson (1910- 1917) reported the common species of Coles County. Weed (1922. 1923) studied the amphibians and reptiles of the Meredosia area, and Blanchard {\92Ab) reported sev- eral new records for Monroe and Randolph counties. Van Cleave (1928) prepared tab- ular keys for the identification of the snakes of Illinois. About 1930 K. P. Schmidt stimulated an interest in the amphibians and reptiles of the Chicago region, and his popular ac- counts of the amphibians (1929, 1930) were followed by a number of papers by Walter L. Necker and others. Schmidt & Necker (1935) published a useful report on the Chicago area species and cited specific locali- ties for all the known amphibians and rep- tiles listed. W. F. Stanley, working for the Illinois Natural History Survey, accumu- lated a considerable number of Illinois snakes through co-operation of the Civilian Conservation Corps. Burt (1928-1931) cited Illinois specimens in several brief pa- pers on the herpetofauna of the Midwest. Goodnight (1937) prepared a key to the salamanders of Illinois, and Cahn (1937) published his excellent monograph of the turtles of the state. Ii.ijNOis Natlrai, Histo?vY SuRvtv Bulletin Vol. 28, Art. 1 About 1940 Cagle beji^an publisbinji the results of his studies on the fauna of south- ern Illinois and in 1941 he published an illustrated key to the amphibians and rep- tiles known for the entire state. This much- needed key was the third paper to treat all amphibian and reptile groups of Illinois. Thirty species of amphibians and 67 of rep- tiles were recorded as occurring in the state. Owens (1941) reported the common spe- cies in Macoupin County, and Lueth (1941) wrote a popular manual of the snakes of Illinois. Peters (1942) listed the species known to occur in Cumberland County, and Cagle (1942fl) published a detailed account of the herpetofauna of Union and Jackson counties. Pope (1944^) summarized the distributional information of the several papers on the amphibians and reptiles of the Chicago area in his extremely useful hand- book on the species occurring in northeast- ern Illinois. Langebartel (1947) reported the snakes known to occur in Hancock County and added Carphophis amoenus vermis to the known fauna of the state. P. W. Smith (1947) analyzed the herpetofauna of east-central Illinois and recorded 50 spe- cies and subspecies for that part of the state. Edgren & Stille (1948) published several papers on the species inhabiting the Chicago region and they added numerous distribu- tional records for northeastern Illinois. Minton & Alinton (1948) cited a number of specimens from southern Illinois, and Grimmer & Langebartel (1948) reported Rurycea longicauila melanopleura as a state record for Illinois. P. W. Smith (1948) added Plethodon dorsalis and Gastrophryne carolinensis to the known amphibian fauna and confirmed the suspected occurrence of Desinognnthus ftiscus and Scaphiopus hol- hrooki in Illinois. Smith & Burger (1950) reported the first Illinois specimens of Masticophis flagellum, Elaphe guttata, and Tantilla gracilis, and P. W. Smith (1951) described and named Pseudacris streckeri illinoensis and Kino- sternon flavescens spooneri from western Illinois sand prairies. Bennett (1953) added Cemophora doliata to the known snake fauna. P. W. Smith (1953-1957) summar- ized the known distribution in Illinois of Tropidoclonion lineatum and Hyla avivoca; resurrected the name semifasciata for the population of garter snakes occurring in northeastern Illinois: called attention to a south-north clinal reduction in number of body blotches in snakes, a reduction that presumably is a manifestation of Gloger's Rule; and offered a reconstruction of the post-Wisconsin biogeography of Illinois and neighboring states. Smith & Minton (1957) summarized distributional data for Indiana and Illinois and discussed postglacial dis- persal patterns of amphibians and reptiles. ILLINOIS AS A HERPETOFAUNAL REGION The natural areas of Illinois have had a great amount of human alteration within historic times. Many natural habitats have been transformed into cities or cornfields, others greatly reduced in extent, and still others variously modified from their orig- inal condition. In a sense, all of the habitats are relictual, separated from each other by expanses of cultivated fields. The state could be described as a great corn desert containing remnants of many habitat types. Although herpetological collecting is rela- tively poor in most parts of the state, Illi- nois has an unusually large number of re- corded species and subspecies. The rich- ness of its fauna, from the standpoint of numbers of species, is due to its great length from north to south, its ecologically inter- mediate position, its biogeographical history, and its variety of habitat types. Physical Features Illinois, fig. 1, has a maximum length of approximately 385 miles from north to south and a maximum width of 215 miles from east to west. The state, which contains 56,400 square miles, is moderately well drained by rivers flowing into the Mississippi on the west, the Ohio on the south, and the Wa- bash on the east. The greater part of the state is a flat or rolling plain. The eleva- tion, which averages about 600 feet for the entire state, decreases from north to south. The lowest part of Illinois is in Alexander County, at the southern tip, approximately 300 feet above sea level. The Shawnee Hills, which extend across the state near the southern tip, the unglaciated area in extreme northwestern Illinois, and a nar- row belt of rock bluff and dissected hills along the Mississippi River have moderate relief; the first two areas have elevations slightly over 1,000 feet. iber, 1961 Smith : Amphibians and Reptiles of Illinois A. ROCK RIVER B. GREEN RIVER C. FOX RIVER D. DES PLAINES RIVER E. KANKAKEE RIVER F. VERMILION RIVER G. SANGAMON RIVER H. ILLINOIS RIVER I. MISSISSIPPI RIVER J. KASKASKIA RIVER K. EMBARRASS RIVER L. WABASH RIVER M LITTLE V\/ABASH RIVER N. BIG MUDDY RIVER O. CACHE RIVER P. OHIO RIVER Q. SALINE RIVER Fig. 1.—Map of Illinois showing counties and major rive Illinois Natural History Survey Bulletin Vol. 28, Art. 1 T[LL_PLAINS_SECTION. GREAT LAKt v/w^ONSInTi "T" 'T'TT !F"n SECTION ,.^^ \HECTI0N)I Rock- llv«r j( -o CENTRAL ^^•^—1'~ "^—J---J\—-T^o-- Hillj Country /-J/ H t-^ _ . V ° !/chlcogo , ! ? L.-.-i-U)V'N i L-. Konkakce LOWLAND PROVINCE Plain .-r.^.-S--% r-L ^ -1._ i ._.H-^""— ^ hri ! £>.=M '9/n VV-^T-^-rH -^- I ! Plain: ' I- . r -—-v-H " L.. I .r( r--rf iLUNOts STAre eeoLoe/CAL suitver Mt. Vernon I ) 1 -i_ ! ;"'""tCENTRAL i H.ll) "^CountVy /lowland ^— i ! L._J province r . SH7wNEE!mLU-Slnw^'^^ 'section jJ^-'LOW \ / plateaus province coastal plain province Fig. 2.—Physiographic divisions of Illinois. From Leighton, Ekblaw, & Horberg (1948] Sovember, 1961 Smith : Amphibians and Reptiles o? Illinois The present topography reflects the Pleis- ocene history of the area. Although four tages of glacial advance, separated by inter- ;lacial periods of milder climate, are known o have occurred, the Illinoian lobe (over 100,000 years ago) and the Wisconsin lobe (about 20,000 years ago) are largely re- sponsible for the physiography of Illinois. Much of the northern half of the state is covered with thick deposits of relatively re- Freeport Mean Jan. temp. Mean July temp. Mean annual temp. Mean inches snowfall Frostfree days Annual precipitation 19.7 73.7 47.8 32.1 -160 -34 nx Chicago Mean Jan. temp. 24.6 Mean July temp. 73.0 Mean annual temp. 49.4 Mean inches snowfall 33.3 Frostfree days 160-170 Annual precipitation -34 Fig. 3.—Temperature and precipitation data for five Illinois cities. Data from Page (1949). 8 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 cent Wisconsin till. Superimposed on this in gradually increasing depths to the west are deposits of fertile loess. The flat or rolling plains are marked by series of low- concentrically arranged recessional mo- raines, the southernmost of which is the Shelbyville terminal moraine. The glaciated area south of the Shelby- ville moraine has deposits of older, less fer- tile Illinoian till, and moraines are no longer evident. This region has slightly more re- lief than the area north of the Shelbyville moraine, since the processes of erosion have had a longer time to cut valleys. Rock outcrops are prominent features in the landscape in the Shawnee Hills, in the northwestern unglaciated area, and along the large rivers. Rock strata appear above the glacial till in small areas throughout the state, particularly In extreme eastern and western Illinois. The physiographic divisions of Illinois have been described by Leighton et al. (1948). These divisions are shown in fig. 2. Climatic Features Illinois has a continental climate. The isotherms are in general from east to west and indicate an increase in temperature from north to south. The difference in av- erage temperatures between northern and southern Illinois is much less marked in summer than in winter. The isohyets are somewhat less regular than the isotherms; in general, the annual precipitation increases from north to south. Union County re- ceives a greater amount of rainfall than any other county. The influence of latitude on temperature and the geographic variation in precipita- tion are illustrated by statistics taken over a period of several years at scattered Illi- nois localities, fig. 3. Biotic Provinces According to the Merriam (1894) classi- fication, Illinois contains three life zones. Extreme northern Illinois is regarded as Transition, extreme southern Illinois as Austroriparian, and the area in between as Carolinian. According to the Shantz & Zon (1924) classification, Illinois includes four vegetation types. Extreme northeastern Illi- nois is classed as Marsh Grass, most of northern and central Illinois as Tall Grass Prairie, southern Illinois, except for the lowermost tip, as Oak-Hickory Forest, and extreme southern Illinois as River Bottom Forest (cypress-tupelo-red gum). According to Dice (1943), the state has three biotic provinces. The northern half is regarded as Illinoian; the southern half, except for the southern tip, as Carolinian; and extreme southern Illinois as Austroriparian. Ken- deigh (1951) indicates the presence of two biomes in Illinois. The northern half is pre- dominantly Grassland Biome, the southern half Temperate Deciduous Forest Biome. The forest is subdivided into three asso- ciations, the largest of which is an ecotone between oak-hickory forest and humid prai- rie. The animal communities are regarded as a Deciduous Forest Biociation, a Decid- uous Forest-Edge Biocies, and a Prairie Biociation. All of these classifications are based on the vegetation or biota of the entire conti- nent, and the boundaries are necessarily in- exact in any minute geographical area. It is noteworthy that all of them indicate that the southern tip of this state is ecologically distinct, and most of them indicate in cen- tral Illinois an ecological break that roughly coincides with the southern limit of Wiscon- sin glaciation. A detailed classification of zones in Illi- nois is expressed by Vestal (1931), who based his divisions upon forest types. In many parts of the state, the distribution of the herpetofauna parallels remarkably the distribution of these types. Vestal's map of Illinois has the following eight divisions: Grand Prairie, Western Division, Jo Da- viess Hills, Mississippi Border, Southern Division, Wabash Border, Ozark Hills, and Tertiary Division, as shown in fig. 4. Herpetofauna! Divisions Although the herpetofaunal distribution tends to coincide with the vegetational divi- sions of Vestal (1931), certain differences exist that should be recognized. The dis- tinctive herpetofaunal divisions are outlined below and shown in fig. 5. I. Prairie or former prairie A. Sand areas B. Grand Prairie (broad sense) C. Outlier prairies II. Forest A. Northeastern Illinois Mesic For- est B. Western Division November, 1961 Smith : Amphibians and Reptiles of iLLixoifi PRAIRIE FOREST Fig. 4.—Vegetation map of Illinois, based on original cover. From Vestal (1931). Forest occupies a relatively small per cent of Illinois at present. Illinois Natural History Survey Bulletin Vol. 28. Art. 1 Prairie W^Si Northeastern Mesic '""" Woodlands Division ^^^^ Woodlands of the Grand Prairie Division Western Division Woodlands Southern Division Woodlands Shawnee Hills Division Austroriparian Division Lower Mississippi Border Division Upper Mississippi Border Division Wabash Border Division Sand Areas Fig. 5.-H.rpe,ofau„al divisions of lUinois J^;jitlllZrttZZlToi^S'T'l -Vefr;ri"s^t'<%ti=e'Vario„ 'i^ri^rz^^:^:!:?.. .»« c„^as.a, P,ai„. S'ovember, 1961 Smith : Amphibians and Reptiles OF Illinois 11 C. Upper Mississippi Border D. Lower Mississippi Border E. Southern Division F. Wabash Border G. Shawnee Hills H. Austroriparian Ninety-four species of amphibians and eptiles occur in Illinois; 25 are state-wide n distribution and 69 occupy limited parts )f the state. The edges of ranges of these j9 have been plotted on one map, fig. 6. It s evident from this map that the ranges of he various species are not uniformly shin- gled over the state but that the range edges )f many tend to coincide. Some areas are elatively uniform ecologically and have ew species reaching the edges of their anges therein. Fig. 6 demonstrates the va- idity of fig. 5. The Sand Areas.—The areas of sand )rairie, Avhich are relatively small, are Fig. 6.—Limits of distribution of the am- phibians and reptiles of Illinois. Each line represents the edge of range of a species. In some areas, lines coincide. chiefly in central and northern Illinois. The largest area extends along the east side of the Illinois River from Woodford County south to northern Scott County. Smaller sand areas occur in Jo Daviess, Carroll, Rock Island, Whiteside, Mercer, and Hen- derson counties, along the east side of the Mississippi River; in Ogle and Lee counties; in Winnebago County; in Kankakee and Iro- quois counties. Small areas of sandy loam occur in Lawrence, La Salle, and Lake coun- ties. A typical sand prairie habitat is shown in fig. 7. The most abundant reptiles of the sand areas are Cnemidophoriis sexUneatus, Pituo- phis melanoleucus sayi. Coluber constrictor flaviventris, Heterodon platyrhinos, Hetero- don nasicus, Terrapene ornata, and Tri- onyx muticus. The distinctive species of amphibians and reptiles are P se udacris streckeri illinoensis, Kinosternon flavescens spooneri, and some of the reptiles mentioned as abundant are also characteristic. A de- pauperate salamander fauna and the local predominance of Bufo woodhousei fowleri over Bufo americanus americanus are also characteristic of this division. The Grand Prairie.—This division as here defined comprises all original prairie, except the sand areas, north of the southern limit of Wisconsin glaciation. It includes accordingly the Grand Prairie (in the strict sense), the Mendota Prairie, and most of the prairies included in Vestal's Western Division. Much of this area has fertile soil, consisting of Wisconsin glacial till and loess, and is now extensively cultivated. It has been characterized by various authors as muck prairie, black-soil prairie, and heavy- soil prairie. The eastern part of the area was formerly dotted with marshes and ponds, most of which have been drained. The most abundant species of amphib- ians and reptiles are Bufo americanus amer- icanus, Thamnophis radix, Elaphe vulpina, Pituophis melanoleucus sayi; probably Sis- trurus catenatus and Emydoidea blandingi were once abundant. The distinctive spe- cies are Matrix kirtlandi, Tropidoclonion lineaium, Opheodrys vernalis, as well as some of the above species indicated as abun- dant. Although none of these is entirely re- stricted to the Grand Prairie, each reaches its greatest abundance in the region. The Outlier Prairies.—These fingers of prairie, which interdigitate with forest in the 12 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 Fig. 7.—A typical sand prairie habitat near Thomson, Whiteside County, Illinois. southern half of Illinois, were included by Vestal in the Southern Division. The north- ern edges of this division are not sharply delimited from the Grand Prairie, and the Springfield District of Vestal is, in fact, eco- tonal in herpetofaunal composition. The soil is rather poor, consisting of clay loam with hardpan a foot or two below the sur- face. The abundant species are Ambystoma texanum, Bufo woodhousei fowleri. Coluber constrictor, Lampropeltis calligaster, and Heterodon platyrhinos. The distinctive spe- cies are Rana areolata and Terrapene or- nate. A few relict colonies of Opheodrys vernalis, Thamnophis radix, Sistrurus cate- natus, and Natrix grahami link this divi- sion with the Grand Prairie. The erratic occurrence of some of these scattered colo- nies is difficult to explain. It seems unlikely that cultivation is solely responsible. Northeastern Illinois Mesic Forest. — The forested moraines and bogs near Lake Michigan contain a relatively rich fauna and are distinctive chiefly because of the occurrence of northern and eastern ele- ments. The abundant species are Ambys- toma marulatum, Rana sylvatica, Lajnpro- peltis triangtilum triangitlum, Storeria oc- cipitomaculata, and Natrix septemvittata. The distinctive species are Ambystoma lat- erale, Hemidactylium scutatum, Clemmys guttata, and ThaTJinophis sirtalis semifasci- ata. The Western Division.—The small patches of woodland and gallery forest along rivers and creeks of this division are rather remarkable for their impoverished fauna. Although apparently suitable habitat for many forest species is available, most of the species known in adjacent divisions have not been found in these forested areas. Acris crepitans, Pseudacris triseriata tri- seriata, Rana pipiens pipiens, Rana clami- tans, Thamnophis sirtalis, and Lampropeltis triangulum are moderately common; all arc more abundant in other parts of Illinois. The Upper Mississippi Border.—^This division, which includes the forested bluffs of the Mississippi River and the Jo Daviess County driftless area, is much richer in number of species than the Western Divi- sion. The extensive rock habitat and the continuity of forest along the Mississippi, which serves as an avenue for the north- ward dispersal of southern animals, prob- ably accounts for the large number of spe- cies in this area. The common species are Natrix sipedon, Diadophis punctatus, Elaphe obsoleta, Thamnophis sirtalis, and Rana clamitans. The distinctive species are Eurycea longi- cauda melanopleura, Carphophis amoenus vermis, Diadophis punctatus arnyi, and Rana palustris. The Lower Mississippi Border.—The bluffs and dissected upland along the Mis- sissippi River south of St. Louis have a spe- November, 1961 Smith : Amphibians and Reptiles of Illinois 13 cies composition and an order of relative abundance similar to that of the Upper Mississippi Border. This division has more affinities with the Shawnee Hills, however, and has, in addition, several western and southern elements not found elsewhere in the state. The distinctive species are Tan- tilla gracilis, Masticopliis flagelluni, Elaphe guttata emoryi, and Gastrophryne carolin- e II sis. The Shawnee Hi»ls.—This division is characterized by forested, dissected upland with sheer blufifs and rock outcrops. The streams, usually fast and clear, have rocky bottoms. The area is rich in numbers of species and individuals. The abundant spe- cies are Ambystonia opacum, Plethodon glutinosus, Eiirycea longicauda, Eurycea lu- cifuga, Hyla crucifer, Hyla versicolor, Sce- loporus undulatus, Scincella laterale, Eu- meces spp., Agkistrodon contortrix, Elaphe obsoleta. Coluber constrictor, Thamnophis sauritus, Virginia valeriae, and Terrapene Carolina. The Southern Division.—The forest of this part of Illinois is largely restricted to river bottoms and land generally unsuited for farming. The habitat is similar to that in the Shawnee Hills but has less relief and few rock outcrops and rocky streams. The unintentional creation of parklike savannas by removal of forest provides ideal habitat for forest-edge species. The thickets, brambles, and grass {Andropogon) glades characteristic of this habitat are in- habited by Bufo woodhousei fowleri, Am- bystoma texanum, Heterodon platyrhinos . Coluber constrictor, Opheodrys aestivus, Storeria dekayi, and Natrix sipedon pleu- ralis. The interior of the woodlands contain Eumeces spp., Elaphe obsoleta, Lampropeltis trianguluni syspila, and Hyla versicolor. The Wabash Border.—The area along the Wabash River in southeastern Illinois is similar to the Southern Division, differ- ing primarily in the higher percentage of forested land and the mesic condition of this forest. Characteristic species are Hyla cru- Fig. 8.—A typical austroriparian habitat at Horseshoe Lake, Alexander County, Illinois. 14 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 lifer, Pseudacris triserlata feriarum X tri- seriata, Eitmrces fascia/us, and Terrapene Carolina. Additional characteristic species are Rana sylvatica, Natrix septemvittata, Natrix eryihroyaster neglecta, Eurycea bis- lincata, PletJiodon cinereus, and Elaphe ob- soltta obsoleta X spiloides. The Austroriparian Division.—This area is the Tertiary Division of Vestal (1931), the River-Bottom Forest of Shantz & Zon (1924), and the Austroriparian of earlier authors. It includes the floodplains, fig. 8, of the Mississippi and Ohio rivers in Alexander, Pulaski, Massac, Pope, western Union, and southwestern Jackson counties. A number of species characteristic of the Gulf Coast extend up the Mississippi River valley into this division. Some of these spe- cies are Ambystoma talpoideum, Hyla cin- erea, Hyla avivoca, Scaphiopus holbrooki, Farancia abacura, Natrix cyclopion, Natrix sipedon confttiens, Cemophora doliata, Agkis- trodon piscivorus, Pseudemys fioridana, and Macroclemys temmincki. PRESENTATION OF DATA The bulk of the data for this report was accumulated during the years 1947-1953. Field work was conducted in almost all parts of the state and was particularly in- tensive in the peripheral areas. Records for the approximately 8,000 specimens in the Illinois Natural History Survey collection were supplemented by Illinois records from the major museums of this country, several state institutions, and a few private collec- tions. Complete synonymies for all Illinois spe- cies and subspecies have been compiled, but, because of their bulkiness, they have been replaced by skeleton sjnonymies; the syn- onymy for each species or subspecies con- sists of a chronological outline of all names and name combinations that have been ap- plied to the taxon in the Illinois literature. Only the initial usage of a name or name combination is cited. The remaining biblio- graphic references and lists of published rec- ords from the time of the original descrip- tion through the appearance of the sixth edition of the official check list (Schmidt 1953) are on file at the Illinois Natural His- tory Survey offices, where they are available to anyone interested in the historical as- pects of the Illinois herpetofauna. The initial citation in every case is to the original description, whether or not Illinois is mentioned ; the specific or subspecific name is followed by the name of the describer (with no comma separating them), the year the description was published, the pagina- tion of the description, and, in parentheses, the type locality. A citation other than the original description refers to the first usage of the name in literature pertaining to Illi- nois; a comma follows the specific or sub- specific name, and the reference is abbrevi- ated to author, year, and pagination. The complete reference for each citation is given in the "Literature Cited" section of this paper. If a citation refers to two different en- tities as now understood, the name appears in the synonymies of both forms involved and is followed by "part" in parentheses. When the citation refers to specimens or populations of intergrades between two sub- species, the name ordinarily is included in the synonymies of both forms and is fol- lowed by "I" in parentheses. The various names and name combinations have been carefully evaluated, and most of them have been definitely allocated by judicious use of our present knowledge of distribution. In a few cases some doubt about assignment of the name remains; in each of these cases the name is prefaced by a question mark. In several instances questionable records in the literature have been clarified by exam- ination of the museum specimens on which the records were based. The diagnosis for each species is brief and includes only those characters or com- binations of characters serving to distinguish an animal from related or similar animals. In most instances it is designed merely to supplement the keys. Exceptions are made, however, for those populations which depart in structure, pattern, or color from pub- lished descriptions of the species or sub- species. The section on variation in each species or subspecies includes a discussion of sexual dimorphism, ontogenetic variation, and geo- graphic variation within Illinois. In an ani- mal that occurs in a limited part of the state, individual variation is summarized. It has been assumed, in accordance with present species concepts, that each individual population probably differs to some extent from all others. Extremes and means of 1 November, 1961 Smith: Amphibians and Reptiles of Illinois 15 counts and measurements rather than more detailed statistics are given, since they illus- trate general trends and do not require cor- rection factors for the small size of the sam- ples. In most cases, the grouping of the samples analyzed has been selected with an eye to the probable natural limits of freely interbreeding populations. In some cases, however, the grouping has been dictated by the availability of specimens. The habits section for each species or subspecies is intended only to give a brief summary of behavior, feeding habits, and life history information. Although a wealth of published information is available for some of the species, little of this informa- tion is based on Illinois animals. The in- formation in the present paper, for the most part, pertains only to Illinois populations; geographic variation in habits as well as in morphology is to be expected. The section on distribution of each species or subspecies in Illinois contains a general statement of the range of the animal in the state, an indication of its general abundance, apparent correlations of its range and the environmental features, and the extent of the intergrading areas of subspecies. In the interest of saving space, lists of locality rec- ords and specimens examined have been omitted. This information is on file at the Illinois Natural History Survey offices and can be transcribed for persons wishing to know precise localities and catalog numbers of specimens on which a record is based. Published records believed valid but not substantiated by museum specimens are listed alphabetically by counties and spe- cific localities and followed by references to the literature cited. The accompanying maps indicate graph- ically the known Illinois localities for the various species. Localities from which specimens have been examined are indicated by dots; localities for which there are liter- ature records believed valid, but from which I have not seen specimens, are indicated by hollow circles. The presumed range of a species that does not occur throughout Illi- nois is indicated by parallel hatching and is based in part on the distribution of appar- ently suitable habitat. In the case of a spe- cies having two or more subspecies in Illi- nois, the range of each subspecies is indi- cated by a different pattern, and the inter- grading area by an overlap of these patterns to form a Crosshatch pattern. The map of the United States that accompanies each Illinois map indicates the total known range of a species and in most cases is from Co- nant (1958). SYSTEMATIC ACCOUNT Amphibians and reptiles comprise two distinct classes of vertebrate animals. The modern species are survivors of those groups of vertebrates that first colonized land. They are intermediate between the bony fishes of an aquatic environment and the birds and mammals of a terrestrial environ- ment. In general, the organ systems of am- phibians reflect the series of profound changes necessary for the transition from water- to air-breathing, from swimming to locomotion on land, and from an environ- ment emphasizing the need for water excre- tion to one requiring water conservation. The organ systems of reptiles reflect the various modifications which made their land colonization more successful than that of the amphibians. In addition to gross differences in organ systems, there are numerous other anatomi- cal features distinguishing the amphibians from the reptiles. These differences are treated in any standard textbook of com- parative anatomy. The most obvious fea- tures distinguishing the amphibians from the reptiles are as follows: Skin without scales or plates ; front feet with 4 or fewer toes; toes without claws; egg« with gelatinous coating; typically with an aquatic larval stage Class Amphibia, p. 21 Skin with scales or plates, or both ; front feet, if present, with 5 toes; some toes with claws; eggs with a tough membranous or limy covering; no larval stage Class Reptilia, p. 113 Check List of the Amphibians and Reptiles of Illinois Class Amphibia Subclass Lepospondyli Order Caudata Family Cryptobranchidae Cryptobranchus alleganiensis al- leganiensis (Daudin) Family Ambystomatidae Ambystoma laterale Hallowell Ambystoma Ttiaculatum (Shaw) 16 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 Ambystoma opacuni (Graven- horst) Ambystoma talpoideum (Hol- brook) Ambystoma ttxanum (Matthes) Ambystoma tigrinum tigrinum (Green) Family Salamandridae Notophtlialmus viridescens lou- isianensis WolterstorfF Family Plethodontidae Eurycea bislineata rivicola Mit- tleman Eurycea long'rauda Irtni/icduda (Green) Eurycea longicauda melanopleu- ra (Cope) Eurycea lucifuga Rafinesque Hemidactylium scutatum (Schle- gel) Plethodon cinereus cinereus (Green) Plethodon dorsalis Cope Plethodon glutinosus glntinosus (Green) Desmognathus fuscits conanti Rossman Family Proteidae Necturus maculosus 7naculosus (Rafinesque) Family Sirenidae Siren intermedia nettingi Goin Subclass Apsidospondyli Order Salientia Family Pelobatidae Scaphiopus holbrooki (Harlan) Family Bufonidae Bufo americanus americanus Holbrook Bufo americanus charlesmithi Bragg Bufo woodhousei fowleri Hinck- ley Family Hylidae Acris crepitans blanchardi Harper Pseudacris triseriata feriarum (Baird) Pseudacris triseriata triseriata (Wied) Pseudacris streckeri illinoensis Smith Hyla avivoca avivoca Viosca Hyla cinerea (Schneider) Hyla crucifer crucifer Wied Hyla versicolor versicolor Le Conte Family Ranidae Rana areolata circulosa Rice & Davis Rana catesbeiana Shaw , Rana clamitans melanota (Rafinesque) Rana palustris Le Conte Rana pipiens piptens Schreber Rana piptens sphenocephala Cope Rana sylvatica sylvatica Le Conte Rana sylvatica cantabrigensis Baird Family Microhylidae Gastrophryne carolinensis caro- linensis (Holbrook) Class Reptilia Subclass Anapsida Order Testudines Family Chelydridae Chelydra serpentina serpentina (Linnaeus) Macroclemys temmincki (Troost) Family Kinosternidae Sternothaerus odoratus (Latreille) Kinosternon flavescens spooneri Smith Kinosternon subrubrum subru- brum (Lacepede) X hippocre- pis Gray Family Testiidinidae Clemmys guttata (Schneider) Emydoidea blandingi (Hol- brook) Terrapene Carolina Carolina (Linnaeus) Terrapene ornata ornata (Agassiz) Chrysemys picta marginata Agassiz Chrysemys picta belli (Gray) Pseudemys scripta elegans (Wied) Pseudemys floridana hoyi (Agas- siz) X concinna hieroglyphica (Holbrook) Graptemys geographica (Le - Sueur) a Graptemys pseudogeographica (Gray) I November, 1961 Smith : Amphibians and Reptiles of Illinois 17 Family Trionychidae Trionyx muticus muticus Le Sueur Trionyx spinifer spinifer Le Sueur Subclass Lepidosauria Order Squamata Suborder Sauria Family Iguanidae Sceloporus undulatus hyacin- thinus (Green) Family Anguidae Ophisaurus attenuatus attenua- tus (Cope) Family Teiidae Cnemidophorus sexVuieattis sex- lineatus (Linnaeus) Family Scincidae Scincella laterale (Say) Eumeces fasciattts (Linnaeus) Eumeces laticeps (Schneider) Suborder Serpentes Family Colubridae Carphophis amoenus helenae (Kennicott) Carphophis amoenus vermis (Kennicott) Farancia abacura reinwardti Schlegel Diadophis piinctatus arnyi Kennicott Diadophis piinctatus edwardsi (Merrem) Diadophis punctatus stictogenys Cope Heterodon nasicus nasirus Baird & Girard Heterodon platyrhinos Latreille Opiieodrys aestivus (Linnaeus) Opheodrys vernalis hlancliardi Grobman Coluber constrictor fiaviventris Say Masticophis flagellum flagellum (Shaw) Elaphe guttata emoryi (Baird & Girard) Elaphe obsoleta obsoleta (Say) Elaphe obsoleta spiloides Du- meril, Bibron, & Dumeril Elaphe vtilpina vulpina (Baird & Girard) Pituophis melanoleucus sayi (Schlegel) Lampropeltis caUigastcr cnlli- gaster (Harlan) Lampropeltis getulus holbrtioki Stejneger Lampropeltis getulus niger (Yarrow) Lampropeltis triangulum trian- gulum (Lacepede) Lampropeltis triangulum syspila (Cope) Cemophora doliata (Linnaeus) Thamnophis sauritus proximus (Say) Thamnophis radix radix (Baird & Girard) Thamnophis sirtalis sirtalis (Linnaeus) Thamnophis sirtalis semifasci- ata (Cope) Tropidoclonion lineatum linea- tum (Hallowell) Virginia valeriae elegans (Ken- nicott) Storeria dekayi ivrightorum Trapido Storeria occipitomaculata occipi- tomaculata (Storer) Natrix cyclopion cyclopion (Dumeril & Bibron) Natrix erythrogaster fiavigaster Conant Natrix erythrogaster neglecta Conant Natrix grahami (Baird & Gir- ard) Natrix kirtlandi (Kennicott) Natrix rhombifera rhombifera (Hallowell) Natrix septemvittata (Say) Natrix sipedon sipedon (Lin- naeus) Natrix sipedon pleuralis Cope Natrix sipedon confluens Blanchard Tantilla gracilis hallowelli Cope Family Crotalidae Agkistrodon piscivorus leucos- tomus (Troost) Agkistrodon contortrix mokeson (Daudin) Sistrurus catenatus catenatus (Rafinesque) Crotalus horridus horridus Linnaeus The preceding check list contains 109 named species and subspecies of amphibians and reptiles known in Illinois. The numer- 18 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 Table 1.—Number of species and subspecies in the herpetofauna of Illinois and some other states. Groip November, 1961 Smith: Amphibiaxs and Reptiles of Illinois 19 Gleason (1907), Cagle (1941), Brown (1950), and Schmidt (1953). The early references to the species in the state were probably based on misidentified specimens of related hylids; the later references may have been based on the assumption that squirella occurred in extreme southern Illi- nois in view of its alleged occurrence in western Kentucky. Recently, at the sugges- tion of Roger Conant and through the cour- tesy of Roger Barbour, I examined the Ken- tucky specimens and found them to be aber- rant specimens of Pseudacris triseriata feri- arum. The nearest valid record of H. squir- ella is approximately 300 miles south of Illinois, and the species should therefore be dropped from the list of Illinois amphibians. Sternothaerus carinatus (Gray).—This Gulf Coast species was reported as occur- ring in Illinois by Davis & Rice (1883fl, 1883^), Jordan (1888), H. Garman (1892), and Hay (1892^), but it was correctly de- leted from the list of Illinois turtles by Cahn (1937). Terrapene Carolina triunguis (Agassiz). —The evidence for deleting this subspecies is presented in the discussion of geographic variation of T. c. Carolina on page 135. CJirysemys picta dorsalis Agassiz.—The evidence for deleting this subspecies is pre- sented in the discussion of geographic vari- ation of C. p. marginata on page 140. Chrysemys picta picta (Schneider). — Yarrow (1882a) recorded the eastern painted turtle from Mount Carmel, and H. Garman (1890) questioned the record. Cahn (1937) deleted the subspecies from the Illinois list. Clemrnys insculpta (Le Conte).—The be- lief that the wood turtle might be found in northern Illinois was expressed by Stej- neger & Barbour (1933, 1939, 1943), Dit- mars (1936), Cahn (1937), Logier (1939), Breckenridge (1944), and McCauley (1945). Cahn summarized the possibility that the species might occur in this state, pointing out that a record is extant from a locality only 11 miles north of the Wiscon- sin-Illinois state line. There is, however, some question about this record, and C. in- sculpta should be dropped from the known turtle fauna of Illinois until additional evi- dence is available. Trionyx ferox ferox (Schneider).—Ken- nicott (1855) included this southern turtle in his Cook County list, but his specimen undoubtedly belonged to T. spinifer spini- fer, as Schmidt & Necker (1935) pointed out. H. Garman (1892) presumed that ferox might occur in the Ohio and Missis- sippi rivers, but Neill (1951a) showed that ferox is unknown north of southern Geor- gia. Phrynosoma cornutum (Harlan).—Dit- mars (1907) included western Illinois in the range of the Texas horned lizard, al- though his inclusion was presumably based on nothing more than a guess. Several specimens of this lizard have been found in Illinois, most of them in urban areas, but these are almost certainly irregulars in the fauna, inasmuch as P. cornutum is one of the most popular of reptile pets. Carphophis amoenus amoenus (Say). — The references to Carphophiops amoenus in Illinois by Garman (1892), Cope (1900), Jordan (1929), and Ditmars (1929) were based on atypical or misidentified specimens of the subspecies helenae. These references antedate the present subspecies concepts; amoenus is now known to be replaced by helenae west of the Appalachians. Abastor erythrogrammus (Holbrook). — Robert Ridgway was apparently responsible for inclusion of Illinois in the range of the rainbow snake by Jordan (1878, 1888), Cope (1877, 1895a), S. Garman (1883, 1884), Davis & Rice (1883a), and H. Gar- man (1892). There is no other evidence that the species occurs north of Louisiana in the Mississippi River valley. Heterodon simus (Linnaeus).—The re- ports of the southern hognosed snake in Illinois bv Davis & Rice (1883a), Jordan (1888), H. Garman (1892), Hart & Glea- son (1907), and Van Cleave (1928) un- doubtedly refer to H. nasicus. The species simus is now known to occur no closer to Illinois than coastal North Carolina. Lampropeltis calligaster rhombomaculata (Holbrook).—The reports of the mole snake in Illinois by Davis & Rice (1883a, 1883^), H. Garman (1892), and Jordan (1888) undoubtedly refer to misidentified specimens of L. c. calligaster. The latter subspecies replaces L. c. rhombomaculata in the areas north and west of eastern Mis- sissippi. Lampropeltis triangulum elapsoides (Hol- brook).—The Illinois references to this southern snake (Davis & Rice 1883a, \S83b; Jordan 1888; H. Garman 1892) 20 Illinois Natural History Survey Bulletin Vol. 28. Art. 1 belong in the synonymy of L. t. syspila. L. t. flapsoides occurs no closer to Illinois than eastern Kentucky. Coluber constrictor constrictor Linnaeus. —The evidence for deleting this subspecies is presented in the introductory remarks con- cerning subspecies of Coluber constrictor on page 196. FAaphe guttata guttata (Linnaeus).—The corn snake was reported as occurring in "southern Illinois" or at Mount Carmel by Yarrow (1882a), Davis & Rice (1883^), Hay (1887A, 1892a), H. Garman (1892). Cope (1900), Myers (1926), Van Cleave (1928), Minton (1944), and Perkins (1949). All of these authors, however, merely presumed that the corn snake might occur in the state, or cited Yarrow (1882^) as authority for inclusion of Illinois in the range of the species. Schmidt & Xecker (1935) deleted the species from the list of Chicago area reptiles. Of the generally ac- cepted records of E. g. guttata, the nearest to Illinois is for specimens from Edmonson County, Kentucky. Although the existence of a population of this species in southeast- ern Illinois is possible, it is much more likely that early authors confused this snake with the superficially similar Lampropeltis calligaster. Virginia striatula (Linnaeus).— The brown snake was credited to Illinois bv Cope (1877), Davis & Rice (1883«, 1883*)', H. Garman (1892), Van Cleave (1928), Cagle (1942fl), and Ditmars (1945). Cagle (1941) noted that Illinois specimens were lacking, but he believed the species might occur in southwestern Illinois. Mar- lin Perkins of the Chicago Lincoln Park Zoo stated that he saw the species near Murphysboro, and his recollection was cited by Ditmars and Cagle. His visual record is highly questionable, however, in- asmuch as V. striatula can be distinguished with certainty from I', valeriae only with the aid of a magnifying lens. Accordingly, it seems advisable to delete this species from the list of Illinois snakes. Virginia valeriae valeriae (Baird & Gir- ard).—Yarrow (1882^), Hay (1892a), and Cope (1900) undoubtedly misidentified specimens of /'. v. elegans when they re- corded the nominate subspecies in Illinois. ]\ V. valeriae does not occur west of south- ern Ohio, being replaced by the subspecies elegans in the Midwest. Thamnophis elegans hammondi (Kenni- cott).—Wright & Wright (1952) men- tioned that an old record for "Tropidonotus hammondi" in Illinois had been published. They listed this among the problematical records, and I have been unable to find the original paper on which the record is based. There can be no doubt that either T. sirtalisl or T. radix was mistaken for hammondi, which is a coastal California subspecies. Thamnophis elegans vagrans (Baird & Girard).—The Rocky Mountain garter snake was included in Illinois reports by Davis & Rice (1883a), H. Garman (1892), and Wright & Wright (1952). Undoubt- edly, a specimen of T. sirtalis was misiden- tified as T. e. vagrans. Thamnophis butleri (Cope).—Ditmars (1922) noted that Butler's garter snake oc- curred in eastern Illinois. Davis (1932), having examined the large series of Tham- nophis in the Chicago museums, pointed out that there were no Illinois records of butleri. Necker (1938), Ditmars (1939), Cagle (1941), A. G. Smith (1951), and Wright & Wright (1952) nevertheless pre- sumed that the species was to be found in northeastern Illinois. Conant (1950) pointed out T. butleri was unknown in Illinois, al- though colonies exist in Wisconsin and southern Michigan. In view of the hun- dreds of garter snakes available from the Chicago area and the absence of specimens of T. butleri, I believe the species should be dropped from the list of Illinois reptiles. Thamnophis sauritus sauritus (Lin- naeus).—The evidence for deleting this subspecies is presented in the discussion of geographic variation of T. s. proximus on page 226. Thamnophis sirtalis parietalis (Say). — The evidence for deleting this subspecies is ' given under the discussion of geographic variation of T. s. sirtalis on pages 231—2. Natrix rigida (Say).—Davis & Rice (1883a), Jordan (1888), and Wright & Wright (1952) mentioned Illinois in defin- ing the range of this Gulf Coast water snake. It is almost certain that Davis & Rice misidentified a specimen of N. gra- hami or "N . septemvittata and that, on their authority, other authors listed the species as occurring in the state. Natrix sipedon fasciata (Linnaeus). — The inclusion of this name in the Illinois literature is probably due to the confusion in November, 1961 Smith: Amphibians anb Reptiles of Illinois 21 the nomenclature of A^ sipedon races rather than to misidentification of specimens. Hur- ter (1911), Loding (1922), and Haltom (1931) apparently applied the name fasciata to the race now known as conftuens; Davis & Rice (1883fl), H. Garman (1892), Gaige (1914), and Wright & Wright (1952) to the race now known as pleuralis. N. s. fas- ciata, as now understood, does not occur north of Louisiana in the Mississippi valley. Micrurus fulvius subsp.—Ditmars (1939) and Cagle (1941) recorded the coral snake as occurring as far north as southern Illi- nois. Pope (1944a) excluded Illinois from the range of the species. Perkins (1949) noted that a dubious record of M. f. fulvius exists from southern Illinois. Although there are old records for the coral snake in Ohio and Indiana, Conant (1951) and Link (1951) have shown rather conclusively that these specimens were probably transported from more southern localities. One specimen (SIU 440) labeled "Cairo" is in the collection of Southern Illinois Uni- versity. This example, a female with a total length of 513 mm., has 15 scale rows, 209 ventrals, 40 caudals, 7 upper and lower labials per side, 11 black body bands, and 3 black tail rings. No other data accompany the specimen except "Cairo" on an outside label. Since the nearest unquestionable rec- ord for the species is in central Arkansas, I am inclined to believe that the specimen was probably collected farther south and subsequently placed in a bottle which hap- pened to bear the label "Cairo." Agkistrodon contortrix contortrix (Lin- naeus).—The evidence for deleting this sub- species is presented in the discussion of geo- graphic variation of J. c. mokeson on pages 26^7. Sistrurus miliarius streckeri Gloyd.—A. E. Brown (1901) expressed the possibility that the pigmy rattlesnake might extend eastward to southwestern Illinois. Until the species is found in Illinois or at least adjacent localit es in Missouri, it should not be included in the Illinois fauna. Crotalus horridus atricaudatus Latreille. —The evidence for deleting this subspecies is presented in the discussion of geographic variation of C. h. horridus on page 271. In addition to species credited to Illinois in the literature, I have seen specimens of such obvious waifs as the caiman and py- thon. Glass AMPHIBIA Salamanders and Frogs Amphibians probably were derived from Crossopterygian fishes early in the Devo- nian Age. They share with the Crossoptery- gians a great many anatomical similarities; also, they possess numerous distinguishing characters, most of which are adaptations for a life on land rather than in the water. There are relatively few modern amphib- ians. The fossil record is sufficiently good to allow recognition of about 10 extinct orders in addition to the three living orders. Two subclasses of amphibians are recog- nized, the Lepospondyli and the Apsido- spondyli. The subclass Lepospondyli con- tains two living orders, the Gymnophiona (caecilians of the tropics) and the Caudata (salamanders). The subclass Apsidospon- dyli contains one modern order, the Sali- entia (frogs and toads). The relationships of the subclasses, or- ders, suborders, and families represented in Illinois may be summarized in the follow- ing synopsis. Subclass Lepospondyli.—Vertebral cen- tra formed directly from mesenchyme; arcualia partially ossified. Order Caudata.—Tail well devel- oped ; at least two limbs present. Suborder Cryptobranchoidea.— Angulare free; gill slits present; external gills absent. Family Cryptobranchidae.— Large; flattened; wrinkled; free, fleshy flanges on limbs. Suborder Ambystomatoidea.—An- gulare and articulare fused ; gill slits and external gills usually ab- sent in adult stage. Family Ambystomatidae.—Lungs and costal grooves present. Suborder Salamandroidea.—Para- sphenoid teeth present; gills usu- ally absent in adult stage. Family Salamandridae.— Costal grooves and nasolabial grooves absent; vomeropalatine teeth in long, posteriorly diverging series. Suborder Plethodontoidea.—Lungs absent; vomerine and palatine teeth usually present. Family Plethodontidae.—Costal grooves and nasolabial grooves present. 22 Illinois Natural History Survey Bulletin' Vol. 28, Art. 1 Family Am ph iumidae.— Limbs much reduced; toes 3-3 or less. Suborder Mutabilia.— Angulare and prearticulare fused; no maxil- lary bone; pterygoid and palatine teeth in continuous series. Family Proteidae.—Large aquatic salamanders, each with 3 pairs of bushy, red gills; toes 4—+; no nasolabial grooves. Suborder Meantes.—External gills present; no hind limbs. Family Siren idae.—Teeth absent; toes 4—0; margins of jaws with horny sheaths. Subclass Apsidospondyli.—Vertebral cen- tra formed from subdivision of the sclerotome; arcualia cartilaginous. Order Salientia.—Tail typically ab- sent in adult stage; pelvic girdle and enlarged hind limbs modified for jumping. Suborder Ano.mocoela.—Sacral ver- tebrae procoelous, fused to coccyx; eight presacral vertebrae uniform- ly procoelous or amphicoelous. Family Pelobatidae.—Cutting tu- bercle on metatarsus; cranial crests absent. Suborder Procoela.—Sacral and pre- sacral vertebrae uniformly procoe- lous; free urostyle; sacral verte- brae with two condyles. Family Bufonidae.—Teeth ab- sent; Bidder's organ in both sexes. Family Hylidae.—Teeth present; intercalary cartilages and toe pads present. Suborder Diplasiocoela.—Eight ver- tebrae ; sacral vertebrae bicon- cave; presacral vertebrae procoe- lous. Family Ranidae.—Vomerine teeth present; no transverse fold of skin on head; tympanum usually distinct; mouth large. Family Microhylidae.—Vomerine teeth absent; transverse fold of skin behind head; tympanum us- ually invisible; head narrow and mouth small. For ease in identification, the Illinois spe- cies of Amphibia are diagnosed in two large keys, the salamanders (the order Cau- data) in one and the frogs and toads (the order Salientia) in the other. Illinois rep- resentatives of these two orders may be sep- arated as follows: Tail absent in adult stage; hind legs conspicu- ously large and modified for jumping Order Salientia, p. 66 Tail present in adult stage; hind legs not conspicuously large and modified for jump- ing Order Caudata, p. 22 Order CAUDATA Salamanders Nineteen species and subspecies of sala- manders occur in Illinois. Another species, Amphiuvia means Garden, has been in- cluded in the key, as it may eventually be found in the southern part of the state. Most salamanders are terrestrial as adults, fig. 9, and aquatic as larvae, but even the adults are chained to moist habitats by their susceptibility to desiccation. A few salamanders remain in water throughout their lives; these are permanently gilled species, which are termed neotenous. The species occurring in Illinois range in size from the 4-inch Hemidactylium to the 24- inch Cryptobranchus. Most species are noc- turnal; they remain hidden under logs or rocks during the day and come forth to search for food at night, when the relative humidity is higher. Salamanders are preda- tory, feeding chiefly on those invertebrates most readily available. They are rather sluggish in their movements and harmless to man. Their defense when captured con- sists of secreting slime. Some salamanders like lizards, readily lose their tails, whicl they use as means of escape. While a pred- ator is occupied by a detached, wriggling tail, the animal to which the tail belonged may escape by crawling away. A few sala- manders can climb; some can jump by vio- lently uncoiling their bodies. Typically, the seasonal life history of sal- amanders begins with a spring migration to a pond, the males usually preceding the fe- males by a day or so. When the females arrive, a nuptial dance ensues, the males swimming over the females and nudging them from time to time. Males deposit sperm packets known as spermatophores on submerged sticks or vegetation, and the fe- males receive these packets with their cloacal lips, fertilization thus being Internal. The females attach their eggs, each surrounded I ss November, 1961 Smith: Amphibians and Reptiles of Illinois 23 by concentric layers of jelly, to vegetation or debris in the water. In a few days to sev- eral weeks, gilled larvae hatch from the eggs and each larva rather quickly develops four legs. Typically, metamorphosis occurs in a few months. The newly transformed adults are difficult to identify for a few days before the characteristic features of mature adults appear. There are several departures from this typical life cycle. At least one species prac- tices external fertilization, the male deposit- ing sperm over the newly laid eggs in the manner of most fishes. Some species differ in having a fall mating and fall oviposition. It is not certain that in all species the fe- male picks up the entire spermatophore, and it is probable that in some species sperm cells escape from the packet and swim into the vent of the female salamander. In a few species the eggs are laid near a stream or pond, and the hatchlings must crawl to water or wait until the hatching site is flooded by rains. The most striking devia- tion from the typical life cycle is found in the genus Plethodon. In all species of this genus, the larval period is spent within the large eggs, which are deposited in rock crevices or rotten logs on land. At hatching the miniature salamanders are ready for a terrestrial existence. Other variations in life histories are noted in the habits sections of the following species accounts. Eight of the 19 Illinois salamanders are predominantly eastern in distribution, occur- ring in Illinois in the Wabash Border coun- ties and in some cases extending varying distances northward or southward in the Mississippi River valley but circumventing the central prairie. One of the eastern spe- cies, Des7nognathus fuscus, has apparently entered Illinois via the Mississippi River valley, since the nearest record outside Illi- nois is in western Kentucky rather than In- diana. The eight eastern components are Cryptobranchus alleganiensis, Ambystoma maculatum, A. opacum, Eurycea bislineata rivicola, E. longicauda longicauda, Pletho- don dorsalis, P. glutinosus glutinosiis, and Des?nognathus fuscus. Three species, Ambystoma texanum, A. i'lgrinum, and Eurycea longicauda melano- pleura, have ranges centering to the west of Illinois. The first two species are rela- tively wide ranging and they completely traverse Illinois. The third is an Ozark species found only in the extreme western part of the state. Three species, Ambystoma talpoideum, Notophthalmus viridescens louisianensis, and Siren intermedia nettingi, are clearly south- ern animals. The first species extends north- ward along the Mississippi River to the southern tip of the state; Siren extends northward along the major rivers in the southern two-thirds of Illinois; Notophthal- mus occurs as far north as the northern boundary of the state. Three species, Ambystoma laterale, Pleth- odon cinereus cinereus, and Hemidactylium scutatum, northern in distribution, extend southward into Illinois. Two species cannot be classified accord- ing to distribution. Necturus maculosus maculosus has a range in which Illinois is approximately centered. Eurycea lucifuga is unassignable because of the peculiar re- striction of this species to areas containing limestone caves. Key to the Order Caudata (Salamanders) 1. One pair of legs (Sirenidae) • Siren intermedia nettingi Two pairs of legs 2 2. Legs vestigial; 3 or fewer toes; body eel- like (Amphiumidae) Amphiuma means tridactylum Legs well developed; 4 or more toes; body not eel-like 3 3. Four toes on each hind foot 4 Five toes on each hind foot 5 4. Feathery external gills; total length to 18 inches (Proteidae) Necturus maculosus maculosus No external gills; total length less than 4 inches (Plethodontidae) Hemidactylium scutatum 5. Body flattened and longitudinally wrin- kled; one gill slit on each side of neck; toes with free fleshy margins, fig. 10 (Cryptobranchidae) Cryp- tobranchus alleganiensis alleganiensis Body terete, not wrinkled; no gill slits; toes without fleshy flanges 6 6. Costal grooves present, fig. 9 7 Costal grooves absent (Salamandridae) Notophthalmus viridescens louisianensis 7. Nasolabial grooves absent, fig. 11 (Amby- stomatidae) 8 Nasolabial grooves present, fig. 12 (Pleth- odontidae) 16 8. Premaxillary teeth in two or more series; neck distinctly wider than head; lingual plicae at oblique angles to median longi- tudinal furrow, fig. 13 Ambystoma texanum 24 Ii.i.iNOis Natural History Survky Bulletin Vol. 28, Art. 1 TAIL COSTAL GROOVES Fig. 9.—Lateral view of a salamander, Ambystoma texanum. Premaxiliary teeth in a single series; head wider than neck; lingual plicae radiate. fig. 14 9 9. Pattern consisting of 4 to 8 silvery white crossbands on dorsum Ambystoma opacnm Pattern not consisting of white cross- bands . 10 10. Body long and slender; toes of adpressed limbs barely overlapping Ambystoma laterale Body stout; toes of adpressed limbs over- lapping by length of longest finger or more 11 11. Groundcolor black or brown, with yellow spots 12 Groundcolor black or brown, without yel- low spots 13 12. Two dorsolateral rows of subequal yellow spots; venter and lower sides brown and unspotted Ambystoma maculatum Irregularly spaced and unequal yellow- spots on back and sides and often on venter . Ambystoma tigrinum tigrintim 13. Head width 23 per cent or more of snout- vent length Ambystoma talpoideum Head width less than 23 per cent of snout- vent length 14 Snout-vent length over 50 mm.; back dark or with brassy flecks Immature Ambystoma tigrinum tigrinum Snout-vent length under 50 mm. ; back dark and with or without flecks 15 Dorsum brown, often with brassy flecks; venter yellow or white Immature Ambystoma maculatum Dorsum black or brown, often with ir- regular white blotches; venter brown Immature Ambystoma opacum Light line from eye to angle of jaw, fig. 15 ; tail triangular in cross section Dcsmognathus fuscus conanti No light line from eye to angle of jaw; tail not triangular in cross section 17 17. Tongue with a central pedicel, free all 14. 15 16 12 Figs. 10-15.—Characteristics of salamand- ers: 10, hind foot of Cryptobranchus allegan- iensis ; 11, face view of Ambystoma opacum; 12, face view of Plcthodon glutinosus ; 13, open- mouth view of Ambystoma texanum; 14, open- mouth view of Ambystoma opacum; 15, head of Dcsmognathus fuscus; c, choana ; c, external naris; I, lingual plica; m, maxillary teeth; n, nasolabial groove; p, premaxiliary teeth; t, tongue ; f , vomerine teeth. A dissecting micro- scope is necessary to see most of these struc- tures. November, 1961 Smith: Amphibians and Reptiles of Illinois 25 around ; groundcolor of dorsum yellow, orange, or red ; venter yellow or white 18 Tongue attached in front, free behind and at sides; groundcolor of dorsum black, brown, or dark gray; venter gray, brown, or black 21 18. Three or more costal folds between ad- pressed limbs; yellow, with brown dor- solateral stripes Eurycea bislineata rwicoUi Two or fewer costal folds between ad- pressed limbs . . 19 19. Body rtd or red-orange, with numerous irregular black spots or dashes scattered over back and sides Eurycea lucifuga Body yellow or orange-yellow, with dis- tinct dorsolateral ;tripes . . 20 20. Sides of tail with many vertical dark bars Eurycea longicauda longicauda Sides of tail dark and without vertical bars Eurycea longicauda melanopleura 21. Costal grooves 15 or fewer; belly uni- form gray or black; back flecked with white; maximum length 180 mm. Plethodon glutinosus glutinosus Costal grooves 16 or more; belly usually mottled; back not spotted with white; maximum length 110 mm. 22 22. Middorsal red, orange, or yellow longi- tudinal stripe 23 No middorsal red, orange, or yellow stripe 25 23. Middorsal stripe zigzag in shape Plethodon dorsalis Middorsal stripe with straight edges 24 24. Costal grooves 16-19, usually 17; tail usually les« than snout-vent length : middorsal stripe usually widest on base of tail Plethodon dorsalis Costal grooves 17-20, usually 19; tail usually longer than snout-vent length ; middorsal stripe widest at mid-body Plethodon cinereus cinereiis 25. Fore legs and breast region with red su'^usion; costal grooves usually 17 or 18; tail usually less than snout-vent length Plethodon dorsalis Fore legs and breast region without red markings; costal grooves usually 18 or 19; tail usually longer than snout-vent length Plethodon cinereus cinereus GRYPTOBRANGHIDAE This family contains two fzenera of large, permanently aquatic salamanders, one of which occurs in the Orient and the other in the eastern United States. Cryptobranchus Leuckart Two subspecies are currently recognized in this monotypic genus. The nominate race, which occurs in extreme eastern Illinois, is widespread in eastern North America; the other race is confined to Ozarkian streams of southeastern Missouri and adja- cent Arkansas. Cryptobranchus alleganiensis alleganiensis (Daudin) Hellbender Salamandra alleganiensis Daudin 1803:231 (type locality: Allegheny Mountains in Virginia). Cryptobranchus alleganiensis, Goodnight 1937: 301. Menopoma alleghcniense, Davis & Rice 1883a: 26. Menopoma alleghaniensis, Brendel 1857:254. Cryptobranchus alleghaniensis, H. Garman 1892:380-1. Diagnosis.—A large aquatic salamander (largest Illinois specimen 440 mm. in total length), fig. 16, with a flattened, much wrin- kled body; 5 toes on each hind foot; loose folds of skin along sides of body and pos- terior margins of legs; 1 gill slit on each side of throat; dark dorsum and slightly less dark venter; without external gills. Variation.—Bishop (1941) found that males of this species were proportionately broader bodied and heavier than females. During the breeding season, the male can be recognized by the swollen ridge sur- rounding the vent. Bishop (1941) believed that transforma- tion occurs in the second year. The newly transformed hellbenders differ from large adults by their more prominent black spots. Four Illinois specimens are extant. One of these, CAS 294, is obviously very old and is accompanied only by the label "south- ern Illinois." It is a female, 225 mm. in snout-vent length; although faded, it still has scattered inky flecks on the dorsum and sides of the tail. Three recent specimens, reported by Stein & Smith (1959), range from 267 to 285 mm. in snout-vent length, 418 to 440 mm. in total length; internar- ial d'stances are 2.0, 2.2, and 3.5 times the spiracular diameters. Two of the three are bluish slate in color; one is dirty brown; two are uniformly colored except for very faint mottling; one is minutely flecked with olive. One (from the Ohio River at Cave in Rock) has rather heavy pigmentation on the lower jaw and a spiracle diameter of only 4.5 mm. and thus approaches the Ozarkian race, Cryptobranchus alleganien- sis bishopi. The others are in every way typical of the nominate subspecies; in dor- 26 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 sal pattern, one of the most reliable subspe- cific characters, all four Illinois specimens are assignable to C. a. alleganlensis. Habits.—The hellbender is permanently aquatic, occurring in fast-running water of rivers and large creeks. Although essen- twisted into a single rosary-like string. Egg masses deposited by several females may be found under one flat rock. The nest site may be attended by a male who aerates the eggs by his movements. Eggs laid in mid- September hatch in mid-November. Dur- Fig. 16.—An adult Cryptobranchus alleganiensis alleganiensis from Harrison County, Indi- ana. (Photo by Isabelle Hunt Conant.) The groundcolor is slate gray; the scattered irregular blotches are black. tially nocturnal, in the spring many of these salamanders are caught on baited hooks during the day. According to Bishop (1941), the lungs of the hellbender are well devel- oped, and captive individuals rise to the sur- face at intervals to gulp air. Hellbenders are ugly in appearance and unpleasant to handle because of their ex- treme sliminess. In food habits they are predatory, eating crustaceans, fishes, mol- lusks, insects, and even their own eggs and young. Bishop (1941), who summarized the life history of C. a. alleganiensis, noted that in the eastern United States breeding aggrega- tions are found in August. Fertilization is external. Approximately 300 to 400 eggs, each of which is about 6 mm. in diameter and surrounded by concentric rings of jelly, are laid in multiple strands, which become ing the 2-year larval period, the young hell- benders, which are gilled, gradually acquire the characteristics of the adults. Illinois Distribution.—^Although C. a. alleganiensis was long presumed by many authors to occur in Illinois waters, actual specimens with locality data were not avail- able until recently. Prior to 1900, the sev- eral references to the species apparently were based on hearsay records of fishermen and on the likelihood that the species oc- curred in the Ohio River down to the mouth of the Wabash River. Blatchley (1899) mentioned a Vigo County, Indiana, speci- men presumably taken in the Wabash River. Stein & Smith (1959) summarized and evaluated all of the reports for Illinois. ' Unquestionable recent records for the Ohio and lower Wabash rivers bordering Illinois indicate that the species occurs in the south- November, 1961 Smith : Amphibians and Reptiles of Illinois 27 Fig. 17.—Distribution of Cryptohranchus al- leganiensis allcganiensis. Hatching indicates the presumed range of the species in Illinois; solid circles indicate localities represented by specimens examined during this study; open circles, localities represented by published records believed to be valid. The lower map depicts the total range of the species in the United States. The Blatchley record referred to on page 26 is not plotted. eastern part of this state and may ascend short distances up the major tributaries of the Ohio and Wabash rivers, fig. 17. Although undocumented by specimens, published records for the following locali- ties are believed valid and are indicated on the distribution map by hollow symbols: L.'^WRENCE County: Lawrenceville, Wa- bash River at Vincennes [Indiana] (Stein & Smith 1959) ; Pulaski County: near Ullin (Stein & Smith 1959). AMBYSTOMATIDAE The family includes three genera in the United States and Canada, two of which are confined to the Pacific Northwest and one of which is transcontinental. Ambystoma Tschudi This North American genus contains 12 species known in the United States and Can- ada; several of these species consist of two or more subspecies. Six species and sub- species occur in Illinois. Ambystoma laterale Hallowell Blue-Spotted Salamander Amblysioma laterale Hallowell 1858:352 (type locality: borders of Lake Superior near Marquette, Michigan). Amblystoma jeffersonianum laterale, Davis & Rice 1883«:26. Ambystoma laterale, Minton 1954:178. Amblystoma jeffersonianum nee Green, Davis & Rice 1883«:26. ?Amblystoma jeffersonianum jeffersonianum nee Green, Jordan 1888:178. Ambystoma jeffersonianum nee Green, Hay 1892^:431. Amblystoma jeffersonianum fuscum, H. Gar- man 1892:372-3. fPlethodon glutinosus nee Green, Shelford 1913:181, 183. Diagnosis.—^A medium-sized, slender, dark brown, black, or blue-black salaman- der (largest Illinois specimen 124 mm. in total length), fig. 18, with discrete pale blue or bluish white flecks on venter, sides, and sometimes on the dorsum; usually 12 or 13 costal grooves, counting impressed lines in groin and axilla; toes of adpressed limbs separated or overlapping by no more than 1 costal fold; head of moderate size, distinctly wider than neck. Variation.—Males of this species pos- sess greatly swollen vents during the breed- ing season. At other times the sexes are difficult to distinguish. Minton (1954) noted that toes of adpressed limbs touch or overlap more frequently in males than in fe- males. The toes of the adpressed limbs touch or overlap by as much as 1 costal fold in speci- mens under 40 mm. in snout-vent length; toe tips vary from an overlap of one-half costal fold to a separation of as much as 3 folds in specimens over 40 mm. Juveniles are relatively larger headed than adults. None of our specimens can be assigned to the related Ambystoma jeffersonianum. In- clusive of the impressed lines in the axilla and the groin, there are 12-12 costal grooves in four specimens, 13-13 in eight specimens, and 14—14 in one specimen. Vomerine teeth 28 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 are usually in 4 series, but sometimes the median pair is fused. Vomerine tooth counts for four specimens range from 27 to 39, averaging 32. Maxillary and premaxillary tooth counts for five specimens range from Stille (1954*) found that in the Chicago region the eggs of this species are laid singly or in small clumps attached to litter on the bottoms of the shallow forest ponds. Egg laying occurs in late March and early April. Fig. 18.—An adult Ambystoma laterale from Lake County, Illinois. The groundcolor is black; vague bluish white flecks are scattered over the lower sides. 76 to 101, averaging 85. Nine of the 14 specimens at hand have discrete light flecks on the sides of their tails as well as on their venters. The tail length ranges from 40.4 to 49.5, averaging 46.5, per cent of the snout-vent length. Habits.—Minton (1954) noted that this species is characteristic of wooded, swampy areas where the soil is sandy and that it may occur in patches of woodland that are sur- rounded by prairie. Like most salamanders, individuals of this species may be found un- der logs or other objects. They are above ground in numbers only during the short breeding season. Their food consists of arthropods and annelids. Hatching occurs approximately a month later, and transformation takes place in late June to mid-August, the time depending on the permanency of the ponds. Reliable in- formation on the life history of the blue- spotted salamander is not available; pub- lished observations concern the related Am- bystoma jeffersonianum or refer to a com- posite of the two species. Illinois Distribution.—This northern species is restricted in Illinois to the north- eastern corner of the state, fig. 19. Yarrow (1882fl), who first reported it from Illinois, considered its range as state-wide. The re- ported occurrence of A. jeffersonianum (equals A. laterale) in southern Illinois is November, 1961 Smith: Amphibians and Reptiles of Illinois 29 Fig. 19.—Distribution of Ambystoma later- ale. Hatching indicates the presumed range of the species in Illinois; solid circles indicate localities represented by specimens examined during this study; open circles, localities rep- resented by published records believed to be valid. The lower map depicts the total range of the species in the United States. in error and actually is based on specimens of the superficially similar A. texanum. The occurrence of the species in the southern half of the state was not challenged by many subsequent authors, probably because of 48 USNM specimens from southern Illi- nois localities cataloged as A. jefferson- ianitm. I have examined these specimens (4096 [38] "S. 111."; 8837, 12050 [3], 12056 [2], 12058-60 Mount Carmel; 8777 Belle- ville) and have found them to be A. tex- anum. Although undocumented by specimens, published records for the following localities are believed valid and are indicated on the distribution map, fig. 19, by hollow sym- bols: Lake County: Beach, Highland Park (Schmidt & Necker 1935). Ambystoma maculatum (Shaw) Spotted Salamander Lacerta maciilata Shaw 1802:304 (type local- ity: Carolina, revised to vicinity of Charles- ton, S. C, by Schmidt 1953). Ambystoma maculatum, Schmidt 1930:10-1. Ambystoma punctata, Kennicott 1855:593. Amblystoma punctatiim, Cope 1868:177. Ambystoma punctatum, O'Donnell 1937:1066-7. Diagnosis.—A large, stout, blue-black, slate-colored, or brownish black salaman- der (largest Illinois specimen 190 mm. in total length), fig. 20, with 2 rows of yel- low-white or orange-yellow, usually round- ed, spots extending from the head well onto the tail; sides and venter slate color to Fig. 20.—A subadult Ambystoma maculatum from Alexander County, Illinois. The ground- color is dark brown or black; discrete yellow spots tend to be arranged in two rows down the back. 30 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 Table 2.—Ontogenetic variation in Illinois Atnbystotna maculattitn, as illustrated by a series from Alexander County. Figures in parentheses are numbers of specimens. Characteristic N ovemDer, 1961 Smith: Amphibians and Reptiles of Illinois 31 summer under logs in the swampy flood- plain forest at Horseshoe Lake in Alexan- der County, where the numerous shallow woodland ponds seemingly afford ideal breeding sites. The spotted salamander re- mains hidden under logs or rocks during the day but may be found wandering about on the forest floor at night. It feeds on arthropods, moUusks, and annelids. Early in the spring spotted salamanders migrate to a woodland pond to congregate and breed. The aggregation performs a sort of nuptial dance, after which the males de- posit spermatophores on sticks and other litter, and the females lay eggs in clusters, usually of several dozen, on debris and vege- tation in the pond. The eggs hatch in less than a month, and the larvae transform be- tween late June and mid-August. Illinois Distribution.—The spotted sala- mander is found in wooded regions in north- ern, eastern, and southern Illinois, fig. 21. Like that of many other eastern forest ani- mals, the range of the species in Illinois is bisected by the Grand Prairie, and popula- tions extend short distances northward from the south and southward from the north in the forested Mississippi River val- ley in western Illinois, but apparently they do not meet. In eastern Illinois the distri- bution is sporadic; permanent woodland ponds are necessary for the occurrence of A. maculatum. A record (UIMNH 1745) from Urbana, which is in a prairie region, is open to some doubt. The general occurrence of this salaman- der from the Gulf Coast to well into Can- ada indicates that temperature is less im- portant as a controlling factor than the pres- ence of mesic forest. Although the forested areas in central Illinois appear to afford suitable habitat, they tend to become dry in late summer; there is no evidence that the species occupies these regions. The pub- lished record for Macoupin County (Owens 1941) is a misidentification based on two CNHM specimens that are actually A. tig- j'iniim. Although undocumented by specimens, published records for the following localities are believed valid; most are indicated on the distribution map by hollow symbols: Cook County: Palos Park (Schmidt & Necker 1935); Northfield (Cope 1868); Du Page County: Wooddale (Stille & Edgren 1948); Union County: Anna (Cagle Fig. 21.— Distribution of Ambystoma macu- latum. Hatching indicates the presumed range of the species in Illinois; solid circles indicate localities represented by specimens examined during this study; open circles, localities rep- resented by published records believed to be valid. The lower map depicts the total range of the species in the United States. 1942a); Union County State Forest (Min- ton & Minton 1948); Wabash County: Mount Carmel (Yarrow 1882^). Ambystoma opacum (Gravenhorst) Marbled Salamander Salamandra opaca Gravenhorst 1807:431 (type locality: New York). Amblystoma opacum, Cope 1868:175. Ambystoma opacum, Hurter 1911:78. Diagnosis.—A medium-sized, stout, black salamander (largest Illinois specimen 121 mm. in total length), fig. 22, with 3 to 8 silvery to grayish white crossbands on the back and 4 to 8 white rings around the tail; dorsal crossbands widened laterally, usually forming a pair of dorsolateral light stripes; head wide; toes of adpressed limbs touching to broadly overlapping. 32 Illinois Natural History Survky Bulletin Vol. 28, Art. 1 Fig. 22.—An adult Amhystoma opaciim from Union County, Illinois. The groundcolor is black; the crossbands are white. Variation.—Males of this species can be distinguished from females by the silvery white crossbands which contrast more sharp- ly against the blue-black groundcolor than do the grayish white bands of the females. In preserved specimens the prominently swollen vent of the male, which is larger than that of the female at all seasons, offers the simplest method of determining the sex. Subadults less than 30 mm. from snout to vent are usually quite unlike adults in pat- tern. The chins and venters are light brownish to light slate color, and the dorsal patterns consist of occasional, poorly de- fined, light blotches on the black ground- color and often white "frosting" on the heads and anterior parts of the bodies. The adpressed limbs of this size group overlap by 2 or 3 costal folds. Specimens 30 to 35 mm. in snout-vent length have essentially the patterns of the adults except that the white crossbands anteriorly are less well defined; the venters are blue-black, and the brown or gray chins show evidence of dark- ening. Specimens 40 to 47 mm. in snout- vent length are proportionately shorter- legged, the adpressed limbs overlapping Ij/' to 1 costal fold. Individuals 48 mm. and larger in snout-vent length vary in the de- gree of overlap of adpressed limbs from 1 costal fold to barely touching. The number of teeth appears to vary with size of the in- dividual. No definite trends in geographic varia- tion within Illinois have been discerned in this species. Individual variation for a sam- ple of 41 specimens from southern and east- ern Illinois is as follows: snout-vent length 32 to 59 mm.; total length 57 to 121 mm.; number of costal grooves 11 to 13 (average 12.1) ; toes of adpressed limbs varying from barely touching to overlapping by 31/ costal folds; number of head and body bands 3 to 8 (average 5.8) ; number of tail rings 4 to 8 (average 6.2) ; number of vomerine teeth 29 to 63 (average 48.0) ; number of pre- maxillary-maxillary teeth 79 to 147 (aver- age 110.0). Habits. — Amhystoma opacum is a wood- land animal, more tolerant of aridity than A. maculatum but apparently less tolerant of low temperatures. Since fall and winter rains insure floodplain pools that last long enough for the transformation of the opacum larvae, a suitable breeding site is probably less important as a limiting factor for this species than for maculatum. The marbled salamander is usually found in wooded hilly regions and can be collected under rocks and logs throughout the sum- mer, sometimes in surprisingly dry situa- tions. In September individual salamanders migrate onto floodplains or into swamps and may be found, often in abundance, under logs or sticks, in heavily shaded areas sub- ject to flooding with the autumnal rains. At Horseshoe Lake, Alexander County, however, the species is extremely abundant throughout the summer, and it appears at least as successful there, in the floodplain swamps, as in upland areas. The food of the marbled salamander con- sists of arthropods, annelids, and mollusks. The courtship of this species is similar to that of the spotted salamander, except that it takes place in the fall rather than in the spring and on land rather than in the water. The males deposit spermatophores in damp places, and the females pick them up with their cloacal lips; then each female November, 1961 Smith : Amphibians and Reptiles OF Illinois 33 deposits approximately 100 eggs in a pro- tected depression. Incubation depends on the length of time before the eggs are inun- Fig. 23.—Distribution of Ambystoma opac- um. Hatching indicates the presumed range of the species in Illinois; solid circles indicate localities represented by specimens examined during this study ; open circles, localities rep- resented by published records believed to be valid. The lower map depicts the total range of the species in the United States. dated, and the hatching time is accordingly variable. Normally, metamorphosis occurs the following June or July. Illinois Distribution. — J. opacum occurs throughout the southern third of Illinois, north along the Indiana-Illinois border probably to Iroquois County, fig. 23, but it appears to be abundant only in the southern fourth of the state. Although undocumented by specimens, published records for the following locali- ties are believed valid and are indicated on the distribution map by hollow symbols: Randolph County: (Hurter 1911); Un- ion County: Cobden (H. Garman 1892); Wabash County: Mount Carmel (Yar- row 1882«). Ambystoma talpoideum (Holbrook) Mo!e Salamander Salamandra talpoidea Holbrook 1838:117, pi. 39 (type locality: sea islands on the border of South Carolina). Amblystoma talpoideum, Cope 1868:173. Ambystoma talpoideum, Hay 1892a: 582. Diagnosis.—A medium-sized, extremely stout, broad-headed salamander (largest Illinois specimen 114 mm. in total length), fig. 24, blue-black or brownish black and usually unmarked above except for a sug- gestion of a longitudinal light stripe on the basal third of the tail; usually 11 costal grooves, counting the impressed line above the hind limbs; head width 22 to 32 per cent of the snout-vent length ; snout-vent length 52 to 58 per cent of total length; toes over- lapping when legs are adpressed ; midven- tral dark stripe usually conspicuous. Table 4.—Ontogenetic variation in Illinois Ambystoma talpoideum. Figures in paren- theses are numbers of specimens. 34 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 Variation.—During the breeding season A mbystoma talpoideum males may be dis- tinguished from females by their swollen vents. I have not been able to find external characters for sexing specimens taken at other seasons. Young individuals are proportionately longer-legged, are wider-headed, are more bluish in color, are more often flecked with gray, and have fewer teeth than adults. These trends are summarized in table 4. Bishop (1941) recorded the maximum length for this species as 97 mm. Four of the 16 Horseshoe Lake, Alexander County, specimens exceed that length, and INHS 5267 is 17 mm. longer than Bishop's largest specimen. Costal groove counts are 11-11 in 14 specimens and 10-10 in two specimens. Variation in dentition and body proportions is shown in table 4. The usual color is blue-gray, but occasionally large specimens are gray-brown. The midventral dark longi- tudinal stripe may be margined on either side by a light area and a secondary dark lateroventral stripe. In most living speci- mens the dorsal portion of the base of the tail is lighter than the rest of the body. Comparative material from South Caro- lina and Louisiana, lent through the cour-' tesy of the late E. B. Chamberlain and of F. R. Cagle, agrees rather closely in most de- tails with Illinois samples. Larvae and adults in the series from South Carolina have a somewhat more prominent ventral pattern, and two of the three Louisiana specimens are at variance with both the Illinois and the South Carolina series in the presence of numerous minute tubercles on the sides of the tails, a condition suggestive of sexual dimorphism. Habits.—The mole salamander is appar- ently similar to the marbled salamander in '^f^:^;^fk^^4^0^. Fig. 24.—Adult Ambystoma talpoideum from Alexander County, Illinois. The groundcolor is dark brown or bluish black; the flecks and often the dorsal crest of the tail are gray. November, 1961 Smith: Amphibians and Reptiles of Illinois 35 Fig. 25.—Distribution of Ambystoma talpoi- deum. Hatching indicates the presumed range of the species in Illinois; solid circles indicate localities represented by specimens examined during this study; the open circle indicates a locality represented by a published record believed to be valid. The lower map depicts the total range of the species as currently known. habits, as the two species have been found together within or under rotten logs in cy- press swamps. Although as many as five specimens of A. talpoideum have been collected in one day at Horseshoe Lake, Alexander County, this species is decidedly less common than A. texanum, opacum, or maculatum. A. talpoideum presumably feeds upon worms, moUusks, and arthro- pods. Life-history data for this species in Illi- nois consist of one observation of a breeding aggregation on November 8. Mosimann & Uzzell (1952) report that in South Caro- lina oviposition occurs in late December and that the 10 to 41 eggs are in compact masses attached to small sticks and grass stems in shallow ponds. The larval stage is of ap- proximately 3 to 4 months duration. Illinois Distribution.—A. talpoideum, an uncommon species, is known at present in Illinois only from Alexander, Jackson, Johnson, and Union counties, fig. 25. I have examined the specimens reported by Cagle (1942fl) from near Anna and 6 miles south- west of Jonesboro (CAS 2512 and 2002) and have found them to be A. texanum. The range of A. talpoideum is yet to be delineated in the Mississippi River valley. Although undocumented by specimens, a published record for the following locality is believed valid and is indicated on the dis- tribution map by a hollow symbol: Alex- ander County: Cairo (Cope 1868). Ambystoma texanum (Matthes) Small-Mouthed Salamander Salamandra tcxana Matthes 1855:266 (type locality: woods near Colorado River and Cummings Creek bottom, Fayette County, Texas ; revised to Rio Colorado bottomland by Schmidt 1953). Ambystoma texanum, Stejneger & Barbour 1933:7. Amblystoma microstomum, Cope 1868:207. Ambystoma microstomum, Hurler 1911:73. Chondrotus microstomus. Cope 1889:101-3. Ambystoma jeffersonianum nee Green, Hay 1892fl:431 (part). Amblystoma jeffersonianum jeffersonianum nee Green, Yarrow 1882a: 50. Amblystoma jeffersonianum platineum nee Cope, Yarrow 1882a: 151. Ambystoma talpoideum nee Holbrook, Cagle 1942rt:175 (part). Diagnosis.—A moderate-sized, slate-col- ored, blue-black, or brownish black sala- mander (largest Illinois specimen 171 mm. in total length), fig. 26, often with lichen- like patches of gray on the sides of the tail and trunk; head narrow and mouth small; gular region conspicuously swollen and wider than greatest width of head; vomer- ine teeth less than 35 and never extending laterally beyond the choanae; maxillary-pre- maxillary teeth usually in 2 or more rows in front, the continuous series containing less than 100 teeth; toes of adpressed limbs touching in subadults or separated by 1 to 4 costal folds in large adults; costal grooves 13 to 15. Variation.—Males of Ambystoma tex- anum have prominently swollen vents dur- ing the breeding season. At all seasons, they are slightly smaller and have longer tails, stouter legs, and wider heads than females. 36 Ii.i.iNOis NATt'RAr, History Survey Bui.i.ktin Vol. 28, Art. 1 Fig. 26.—An adult Ambystoma texanum from Calhoun County, Illinois. The groundcolor is dark brown or black, usually with flecks of gray scattered over the back and sides of the animal. Trends in ontogenetic variation are ex- pressed chiefly by proportions and color and are illustrated by a sample of 22 specimens from Horseshoe Lake, Alexander County, table 5. Geographic variation within Illinois is rather slight. A sample (14 specimens) from the southern tip of the state tends to have smaller individuals, slightly lower tooth counts, and perhaps a higher fre- quency of subadults with distinct gray frost- ing on the dorsum than samples from else- where in the state. In 32 per cent of the 14 southern Illinois specimens a single row of premaxillary teeth is characteristic, whereas in only 6 per cent of 37 specimens from other parts of Illinois is a single row characteristic; 2 or 3 rows are typical. Individual variation for a sample of 51 adults is as follows: snout-vent length 50 to 90 mm.; total length 89 to 155 mm.; number of costal grooves 13 to 15 (average 14.2) ; number of vomerine teeth 18 to 33 (average 25.0) ; number of maxillary-pre- maxillary teeth 58 to 101 (average 75.0). A specimen from McLean County has a bifid tail, an anomaly due to an injury to the tail of the animal. Habits.—The small-mouthed salamander is the most common Ambystoma in the state. It occurs in woodlands, prairie, pastured areas, and even in the intensively cultivated black soils of central Illinois. Adults are en- countered during the summer months in various habitats, such as rocky hillsides, creek beds, and floodplain swamps. They feed on earthworms, slugs, and arthropods. They are nocturnal and fossorial. Table 5.—Ontogenetic variation in Illinois Ambystoma texanum. Figures in parentheses are numbers of specimens. Characteristic Snout-vent length as percentage of total length Toes of adpressed limbs Number of vomerine teeth Number of maxillary-premaxillary teeth Color of venter Snout-Vent Length (mm.) 33-40(9) 41-54 ryi 56-64(6) 55.6-59.5 Touch 15-27 32-73 Light 52.3-57.0 20-26 61-77 Light or d'ark* 49.8-58.0 - m 18-27 61-72 Dark "Preserved specimens of this size group trequently liave sharply bicolored side November, 1961 Smith: Amphibians and Reptiles of Illinois 37 Breeding occurs in late February and throughout March in most of Illinois. Al- most any standing water is apparently suit- able as a breeding site inasmuch as eggs have been observed in ditches, flooded fields, woodland streams, and even in cis- terns. The egg masses, each usually con- sisting of 6 to 30 eggs, are deposited on sticks or vegetation in shallow water. Often eggs and spermatophores are present on the same object. The eggs hatch in a few days, the time depending upon the temperature of the water. The larvae transform Into adults from late May through July. Illinois Distribution.—This species in Illinois occurs north to Henry and Kan- kakee counties, fig. 27, and it is abundant in both woodland and prairie regions. The ap- parent gaps in range in extreme western Illinois and south-central Illinois, as shown Fig. 27.—Distribution of Ambystoma tex- anum. Hatching indicates the presumed range of the species in Illinois; solid circles indicate localities represented by specimens examined during this study; open circles, localities rep- resented by published records believed to be valid. The lower map depicts the total range of the species in the United States. on the map, are probably merely indicative of little field work in these regions during early spring. The absence of records from Adams County, a relatively well-collected area, is puzzling. Although undocumented by specimens, published records for the following locali- ties are believed valid and are indicated on the distribution map by hollow symbols: Jackson County: 3 mi. N Carbondale (Cagle 1942a); Knox County: Galesburg (H. Garman 1892). Ambystoma tigrinum tigrinum (Green) Eastern Tiger Salamander Salamandra tif/iina Green 1825:116 (type locality: near Moorestown, N. J.). Amblystoma tigrinum, Cope 1868:183-4. Ambystoma tigrinum, Hurter 1911:75. Ambystoma tigrinum tigrinum, Dunn 1918:457. ?Ambystoma lurida, Kennicott 1855:593. '?Amblystoma copianum, H. Garman 1892:215. Ambystoma v.aculatum nee Shaw, Owens 1941: 183. ?Ambystoma sp., P. W. Smith 1951:192. Diagnosis.—A rather large, blue-black or brownish black salamander (largest Illi- nois specimen 330 mm. in total length), fig. 28, with irregularly arranged yellow blotches, which are unequal and variously shaped, on the back and sides; costal grooves 12 or 13, sometimes 1 1 or 14; toes of adpressed limbs usually overlapping by 2 or more costal folds; 2 palmar tubercles distinct on freshly preserved specimens. Variation.—In Ambystoma tigrinum tig- rinum, the male differs from the female in having a longer vent and a longer tail. Dur- ing the breeding season, the vent of the male is strongly protuberant. Newly metamorphosed specimens, 50 to 60 mm. from snout to vent, are gray, brown, or olive, often with numerous indistinct dark spots above. Slightly older examples are black above, with a few very small yel- low spots. The pattern of the adult is found in specimens 65 to 75 mm. in snout-vent length. The proportionate length of the legs decreases and the number of teeth in- creases with an increase in size. This vari- ation is summarized in table 6. No well-defined geographic variation within Illinois has been discerned for the eastern tiger salamander, although a dif- ference in size between northern and south- ern specimens is indicated. Fifteen speci- i 38 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 Fig. 28.—An adult Ambystoma tigrinum t'tgrinum from Randolph County, Illinois. The groundcolor is dark brown or black; the yellow spots are irregular in size and arrangement. In some large adults the spots are obscured. mens now available from the southern third of the state are less than 100 mm. from snout to vent, and numerous other exam- ples observed but not preserved were well under this length. Fifteen specimens out of 49 now at hand from the northern half of Illinois exceed 100 mm. in snout-vent length. Pope ( 1944/1) noted that the spe- cies attains its maximum size in the Chi- cago area. The largest specimen known (330 mm. in total length) is from Quincy, Illinois (Langebartel 1946). Lengths and costal groove counts for three samples are given in table 7. There is considerable individual varia- tion in markings. Several of the large in- dividuals are so dark that the spotted pat- terns are barely discernible. Other speci- mens, usually smaller individuals, are boldly marbled with black and yellow. Three large Table 6.—Ontogenetic variation in Illinois Ambystoma tigrinum tigrinum. Figures in parentheses are numbers of specimens. Characteristic November, 1961 Smith : Amphibians AND Reptiles of Illinois 39 specimens from central Illinois are light olive, with black spots and mottling. Habits.—Like the small-mouthed sala- mander, the tiger salamander is nocturnal and fossorial. Accordingly, it is seldom seen unless it is discovered under a log or rock, or within some steep-sided excava- tion. Although this salamander is not abun- dant, it persists in the most disturbed areas, often occurring within cities and in inten- sively cultivated regions. Its food consists of almost any kind of animal it can over- power. Captive salamanders of this spe- cies take earthworms and hamburger read- The number of individuals found abroad late in the fall suggests that a migration to the breeding ponds occurs at that season. Actual breeding, however, occurs early in the spring. Loose clusters of 25 to 100 eggs Fig. 29.—Distribution of Ambystoma tigrin- um. The nominate subspecies occurs through- out the state. Solid circles indicate localities jrepresented by specimens examined during ithis study; open circles, localities represented ;by published records believed to be valid. The lower map depicts the total range of the species in the United States. are attached to objects on the bottom of a pond. The incubation period depends on the temperature, probably averaging about 3 weeks. The larvae grow rapidly and in central Illinois transform in July. At meta- morphosis the larvae of this species are about twice the size of other Ambystoma larvae. Illinois Distribution.—This species is state-wide in distribution, fig. 29, but not abundant except in areas with numerous ponds. In most parts of the state, speci- mens are usually found quite incidentally in basements, deep-sided ditches, submerged meter boxes, or similar pits. Many per- manent ponds, either prairie or woodland, harbor the species; small forest-edge ponds are probably preferred. Although undocumented by specimens, published records for the following locali- ties are believed valid and are indicated on the distribution map by hollow symbols: Cook County: Glenview (Necker 1939c) ; Northfield (Yarrow 1882fl) ; Lake Coun- ty: Waukegan (Necker 1939f ) ; La Salle County: La Salle (Burt 1935); Madison County: (Hurter 1893) ; Mason County: SE Havana (Vestal 1913) ; Men- ard County: Athens (Dunn 1918) ; Mon- roe County: (Hurter 1893); Peoria County: (H. Garman 1892). SALAMANDRIDAE This family includes two genera in the United States and Canada; Notophthalmus occurs east of the Great Plains and Taricha on the West Coast. Notophthalmus Rafinesque Thiee species of this eastern North Amer- ican genus are found in the United States and Canada. The species occurring in Illi- nois consists of several subspecies, but only one of them is found within this state. Notophthalmus viridescens louisianensis (Wolterstorff) Central Newt Diemyctylus viridescens louisianensis Wolter- storflF 1914:383 (type locality: New Orleans, Louisiana). Tritiirus viridescens louisianensis, Schmidt & Necker 1935:62. Diemictylus viridescens louisianensis, Brown 1950:24. 40 Illinois Natural History Survev Bulletin Vol. 28, Art. 1 Notophthalmus miniatus, Kennicott 1855:593. Diemyctyltis miniatus miniatus. Yarrow 1882a: 161. Diemyctylus viridescens miniatus, Hay 1887a: 6. Diemyctylus miniatus, H. Garman 1890:190. Triton Jorsalis net Harlan, Brendel 1857:254. Diemyctylus miniatus viridescens ncc Rafiii- esque, Yarrow 1882a: 161. Diemyctylus viridesccns. Hay 1887^:62. Diemyctylus viridescens viridescens nee Rafin- esque, Cope 1889:212-3. Diemictylus viridescens, Shelford 1913:252. Notophthalmus liridescens viridescens nee Ratinesque, Stejneger &: Harbour 1917:8. Triturus viridescens, Dunn 1918:451. Triturus viridescens viridescens nee Raf-n- esque, Stejneger & Barbour 1923:4. Diagnosis.—A small, stout, aquatic or terrestrial salamander (largest Illinois spec- imen 104 mm. in total length), fig. 30, with- out costal grooves; gular fold poorly de- veloped; cranial crests present; vomeropal- atine teeth in two longitudinal series; dor- sum yellow-brown, olive-green, or red- brown and sharply cut off from yellow of venter and lower sides; scattered black specks above and below ; one or two dorso- lateral rows of small red dots, some of which are not margined with black or at least not completely encircled. The eft, a peculiar terrestrial stage found only in this salamander, differs from adults, which are aquatic, by the absence of fins on the tail, by the absence of secondary sexual charac- ters, and by the presence of a granular or warty skin and darker groundcolor. Variation.—Efts are difficult to sex, but the depth of the pits in the temporal region (hedonic glands) serves to identify at least some males. Adults in breeding ponds are easily sexed. The male possesses greatly en- larged hind legs with horny material on the inner sides, a swollen cloaca, and conspicu- ous dorsal tail crest during the breeding season, in addition to the deep hedonic pits at all times. The female in the breeding season has dark, converging, cornified ridges on the lips of the cloaca. No data on ontogenetic variation are available for the Illinois populations. Most of the speci;r.ens extant are efts. No variation trends within Illinois have been discerned. Ten specimens from north- eastern Illinois vary as follows: snout-vent length 18.8 to 40.2 mm.; total length 39 to 81 mm.; lateral red spots enclosed by black, to 8 per side (average 2.3) ; lateral head Fig. 30.—Adult Notophthalmus viridescens louisianensis, eft stage, from Monroe CountyJ Illinois. The groundcolor above is dark brown or olive-brown; below, jellow. November, 1961 Smith : Amphibians and Reptiles of I llixoii 41 stripe prominent in four specimens, weak in five, absent in one. Fourteen specimens from the southern third of the state exhibit the following variation: snout-vent length 18.2 to 45.2 mm.; total length 35 to 104 mm.; lateral red spots ringed with black, to 9 (average 4.4) ; lateral head stripe prominent in 1 specimen, weak in 13. Habits.—Notophthalmiis viridescens has a rather complicated life history, as adults have both a terrestrial and an aquatic stage. Presumably the subspecies louisian- ensis and viridescens have essentially the same life cycle, which in N. v. viridescens includes an underwater nuptial dance and deposition of spermatophores and eggs in winter or early spring. The eggs, which are attached individually to submerged vegeta- tion, hatch in 2 or 3 weeks, and the larvae transform usually in 2 or 3 months. The transformed newt is rather dry and warty- skinned. It soon loses its gills and fins; it is then called an eft. Because the eft lives under bark, logs, or rocks in typical sala- mander habitats, this is the form most often encountered in Illinois. After 2 or 3 years, the eft is sexually mature and ready for an aquatic existence. It develops fins and skin changes that permit aquatic respiration. The number of tiny efts that have been found in southern Illinois suggests that the initial aquatic stage may be shorter in louisianensis than the 2 or 3 months of viridescens. Newts eat a variety of invertebrates, mollusks be- ing a particularly important item. Illinois Distribution.—The central newt [ probably once occurred throughout the [ state except in the extreme western part. I All recent records are for either the northern I or for the southern fifth of Illinois, fig. 31. I Although records for the central part of I the state are at least 35 years old, there is 1 little reason to doubt them, inasmuch as the central newt is easily recognized. It is ' likely that its occurrence in the area be- I tween extreme northern and southern Illi- , nois was sporadic even before cultivation. The destruction of much of the forest, the ' draining of ponds and marshes, and the lowering of the water table probably accel- I erated the disappearance of this salamander I from central Illinois. j Although undocumented by specimens. ! published records for the following localities are believed valid ; most are indicated on the distribution map by hollow symbols: Coi.es Fig. 31.—Distribution of Notophthalmus 'viridescens. Hatching indicates the presumed range of the subspecies louisianensis in Illinois; solid circles indicate localities represented by specimens examined during this study; open circles, localities represented by published rec- ords believed to be valid. The lower map depicts the total range of the species in the United States. County: Charleston (Hankinson 1917); Hardin County: Cave in Rock (H. Gar- man 1892); Jackson County: Grand Tower (H. Garman 1892); Kane Coun- ty: Geneva (H. Garman 1892); McLean County: Normal (Dunn 1918); Peoria County: Peoria (H. Garman 1892); St. Clair County: Bluff Lake (Hurter 1893) ; Tazewell County: Delavan (H. Gar- man 1892); Union County: Anna (Cagle 1942«). PLETHODONTIDAE Four of the 17 plethodontid genera known from the United States and Canada are represented in Illinois. The family is pri- marily New World in distribution but it is represented in Europe by one genus. 42 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 Eurycea Rafinesque The genus Eurycea includes eight species, most of which have several subspecies. Three species occur within Illinois. Eurycea bislineata rivicola Mittleman Midwest Two-Lined Salamander Eurycea bislineata rivicola Mittleman 1949: 89-96 (type locality: Echo Canyon, McCor- mick's Creek State Park, Owen County, Indiana). Eurycea bislineata bislineata nee Green, Dunn 1926:307. Eurycea bislineata, Jordan 1929:218. Spelerpes bilineatus, Davis & Rice 1883a :27. fDesmot/nathus fusea nee Rafinesque, Hank- inson' 1915:293. fEurycea longicauda ncc Green, Peters 1942: 182. Diagnosis.—A small, slender, yellow or tan salamander (largest Illinois specimen 108 mm. in total length), fig. 32, with a lat- eral brown or black band of variable width extending from the eyes almost to the tail tip; 13 to 15 costal grooves; II/2 to 5 costal folds between the adpressed limbs; 6 to 22 vomerine teeth in two arched series behind and between the choanae; and usually either a median series of dark spots or irregularly spaced dark middorsal flecks. Variation.—According to Bishop (1941), the male of this salamander differs from the female in its larger size, wider head, larger premaxillary teeth, and nasolabial tubercles. The teeth of the female at all times, and of the male from June to September, are said to be bicuspid. Ontogenetic variation is not marked in the Illinois specimens available. The differ- ence in ratio of tail length to total length, noted by Oliver & Bailey (1939) and Mit- tleman (1949), is apparent only when the figures are averaged. In 12 specimens 53 to 75 mm. in total length, the tail length ranges from 51.5 to 57.4 per cent (average 55.1) of the total length. In 12 specimens 85 to 108 mm. in total length, the tail length ranges I Fig. 32.—Adult Eurycea bislineata rivicola from Clark County, Illinois. The groundcolor is tan or yellow; the markings are dark brown. November, 1961 Smith : Amphibians and Reptiles of Illinois 43 Table 8.—Geographic variation in Illinois Eurycea bislineata rivicola. Figures in paren- theses are numbers of specimens. Characteristic 44 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 Fig. 33.—Distribution of Eurycca bislincata. Hatching indicates the presumed range of the subspecies rivicola in Illinois; solid circles indicate localities represented by specimens examined during this study; the open circle indicates a locality represented by a pub- lished record believed to be valid. The lower map depicts the total range of the species in the United States. Eurycea longicauda (Green) Four subspecies of this eastern, stream- inhabiting salamander have been described, but only three are recognized in the present study. Two well-marked subspecies occur within Illinois. Eurycea longicauda longicauda (Green) Long-Tailed Salamander Salamandra longicauda Green 1818:351 (type locality: New Jersey; revised to vicinity of Princeton by Schmidt 1953). Eurycea longicauda, Stejneger & Barbour 1917: 19. Eurycea longicauda longicauda, Cagle 19+1:5. Spelerpes longicaudus, Davis & Rice 1883a :27. Eurycea longicauda pernix Mittleman 1942: 101-5, pi. 20 (along Jimmie Strahl Creek, 2'/2 mi. SE Nashville, Brown County State Park, Indiana). Diagnosis.—A moderate -sized, slender, long-tailed salamander (largest Illinois spec- imen 15.^ mm. in total length), fig. 34, with yellow or orange-yellow groundcolor and brown or black maculations that tend to coalesce on the sides to form lateral bands; sides of tail with vertical dark bars; 13 to 14 costal grooves; toes of adpressed limbs overlapping by 2 costal folds or separated by 2 folds; 12 to 39 vomerine teeth in all; venter immaculate yellow; middorsum plain or more often with numerous dark spots clustered along the midline. Remarks.—The recently described Eury- cea longicauda pernix (Mittleman 1942) of unglaciated Indiana is said to differ from E. I. longicauda by a higher vomerine tooth count, more intense pattern, and greater de- velopment of sexual dimorphism. The holo- type and allotype of pernix have, respec- tively, 27 and 32 vomerine teeth. Only 8.6 per cent of the Illinois specimens, exclusive of E. I. melanopleura intergrades, have 27 or more vomerine teeth. Tooth counts for 11 specimens, which should be referable to pernix on the basis of range (Brown, Jen- nings, Morgan, and Owen counties, Indi- ana), vary from 11 to 26 (average 19.6); similar counts for 6 specimens of E. I. longi- cauda from glaciated Indiana (Parke and Montgomery counties, Indiana) vary from 18 to 28 (average 22.2). Accordingly, a high number of vomerine teeth does not ap- pear to be diagnostic for the population m unglaciated Indiana and Illinois. I can discern no differences in development of secondary sexual characters or pattern which will separate specimens of E. longi- cauda from glaciated Indiana from those of unglaciated areas. There is a tendency for the lower sides of specimens from Mary- land and Pennsylvania to have zones of small, well-separated spots, whereas the lower sides of salamanders from the Great Smoky Mountains and transmontane regions are vermiculate. It seems best to refer the Illinois populations to E. I. longicauda, pend- ing a restudy of the species. Variation.—External sexual differences in this salamander are slight. Bishop (1943) reported that the male has more swollen nasolabial tubercles than the female, a slightly papillose vent, a shorter tail, and toes almost meeting when the limbs are November, 1961 Smith: Amphibians and Reptiles of Illinois 45 Fig. 34.—Adult Eurycea longicaiida longicauda X melanopleiira intergrades from Monroe County, Illinois. The specimen at the left is fairly typical of the nominate subspecies; the speci- men at the right shows some melanopleiira influence in the coalescence of the dark bars on the sides of the tail and the spotting on the dorsum. The groundcolor ranges from lemon yellow to almost orange; the markings are dark brown or black. adpressed. In the female the toes of the ad- pressed limbs are separated by at least 1 costal fold. Subadults are quite different from adults in general appearance, possessing a clear yellow groundcolor, smaller middorsal spots, shorter tails, and fewer vomerine teeth. They usually have prominent dark spots on the anterior part of the chins, whereas adults usually have dusky chins. The onto- genetic variation in proportionate tail length and number of vomerine teeth is summa- rized in table 9. Geographic variation for this subspecies in Illinois is manifested by intergradation with the western race, E. I. melanopleiira. Populations of typical longicauda occur in the Wabash Border counties and through- out the Shawnee Hills. A slight influence of melanopleura is seen in the tendency for the vertical tail bars to coalesce in occasional specimens from western Union and Jackson counties. Samples from other Lower Mis- sissippi Border counties exhibit a gradual replacement, south to north, of longicauda characters by those of melanopleura. The Table 9.—Ontogenetic variation in proportionate tail length and in vomerine tooth counts in Illinois Eurycea longicauda. Figures in parentheses are numbers of specimens. 46 Illinois Natural History Survey Bulletin Vol. 28, Art. I longicauda influence predominates as far north as St. Louis, but north of St. Louis specimens possess characters closer to those of melanopleura. A sinjile specimen (INHS), a juvenile, is known from the Wabash Border Division. This juvenile, 54 mm. in total length, has 14 costal grooves, a combined vomerine count of 16, 1 costal fold between toes of the adpressed limbs, a distinctly barred tail, or outcrops by searching these areas with a flashlight. On rainy days, individuals are sometimes abroad and may be seen on trunks of trees. Normally, however, they remain hidden under rocks or logs or within crev- ices in bluffs during the day. Their food consists predominantly of arthropods. Details of the life history of this subspe- cies are lacking, but they are probably sim- ilar to those of E. hisl'ineata rivicola. Larvae Table. 10.—Geographic variation in Illinois Eurycea longicauda. Figures in parentheses are numbers of specimens. Characteristic November, 1961 Smith: Amphibians and Reptiles of Illinois 47 Eurycea longicauda melanopleura (Cope) Dark-Sided Salamander Spelerpes melanopleurus Cope 1893Z':383 (type locality: Raley's Creek, White River, Mis- souri). Eurycea longicauda pernix X E. I. melano- pleura, Mittleman 19+2:104. Eurycea longicauda melanopleura, Grimmer & Langebartel 1948:224. Diagnosis.—A medium-sized, long-tailed salamander (largest Illinois specimen 130 mm. in total length), fig. 35, related to the subspecies longicauda but differing in having a longitudinal dark band (rather than a se- ries of vertical bars) on each side of the tail, numerous middorsal dark spots scat- tered or arranged in several irregular rows, numerous dark flecks on venter, vellow- green to yellow-brown groundcolor, and larger eyes. Variation.—The males of the dark-sided salamander differ externally from the fe- males in having more swollen snouts and larger nasolabial protuberances, but so many intermediate examples occur that dissection is often the only method by which the sex of an individual can be accurately deter- mined. In this subspecies subadults differ from older specimens less markedly than in Eury- cea longicauda longicauda but in similar characters. Seven specimens, 88 to 114 mm. in total length, have tail lengths 48.0 to 65.5 (average 58.3) per cent of the total length, and six specimens, 115 to 130 mm. in total length, have tail lengths 58.5 to 65.2 (av- erage 60.8) per cent. The groundcolor is us- ually more yellowish in small individuals. :*^W. ^"-s. ^Wftfj Fig. 35.—An adult Eurycea longicauda melanopleura from Adams County, Illinois. The groundcolor ranges from greenish yellow to pure yellow; the markings are black or dark brown. 48 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 but the pattern is essentially the same. Some large specimens are suffused with tine flecks of darker pigment. The Illinois population differs slightly from samples from more western localities in the smaller average number of vomerine teeth, in the poorly developed cirri, and in the usual irregularity of the rows of mid- dorsal dark spots. Nevertheless, the sam- ples from western Illinois are homogeneous, quite distinct from the eastern E. I. lotiffi- cauda, and the differences noted are not due to intergradation with the eastern form. Thirteen specimens from Adams County show the following variation: snout-vent length 32.8 to 54.0 mm.; total length 88 to 133 mm.; 13 costal grooves in 3 speci- Fig. 36.— Distribution of Enrycea long'icauda. Vertical hatching indicates the presumed range of E. I. longicauda; horizontal hatching, the presumed range of E. I. melanoplriira; cross- hatching, the area of intergradation between the two subspecies; solid circles indicate local- ities represented by specimens examined dur- ing this study; open circles, localities repre- sented by published records believed to be valid. The lower map shows the total range of the species in the United States. mens, 14 in 10; toes of adpressed limbs overlapping by 1 costal fold to a separation of 2 folds; combined vomerine tooth counts 14 to 30 (average 20.5) ; cirri present in 1 specimen, nasolabial tubercles in 12; scat- tered ventral flecks in 11 specimens, immac- ulate venters in 2. Intergrades of melanopleura X longicaudi have been discussed under the subspecies loiujicauda. Samples from Pike and Greene counties display some characters of longi- cauda but the melanopleura influence is sol pronounced that these specimens would keyj out as melanopleura. Habits.—This subspecies is similar to thej eastern long-tailed salamander in habitsJ The only difference that seems worthy ofj notice is the more frequent occurrence ofj melanopleura within caves. Illinois Distribution.—This salamandei is common in rocky streams, caves, andl springs of the Mississippi River bluffs andj dissected wooded uplands in Adams anc Pike counties, fig. 36. Intergrading popula- tions occur south to Union County, but onlyl those populations north of St. Louis have! the general appearance of melanopleura.\ The apparent absence of this subspecies north of Adams County may be due to tem- perature intolerance, inasmuch as appar- ently suitable habitat extends well north of the known distribution. The upper Illinois River valley appears to afford suitable habi- tat, but none of the subspecies of E. longi- cauda has been able to ascend this valley^ probably because the lower portion lacks the requisite rocky streams. Although undocumented by specimens, published record for the following locality is believed valid and is indicated on the dis-i tribution map by a hollow symbol: Adams County: 11 mi. S Quincy (Grimmer Langebartel 1948). Eurycea lucifuga Rafinesque Cave Salamander Eurycea lucifuga Rafinesque 1822:3 (typfi locality: caves near Lexington, Kentucky) Stejneger & Barbour 1917:19. ?Spclerpes ruber ncc Latreille, Hay 1887. foiL'leri undoubtedly refers to Table 16.—Geographic variation in four samples of sexually mature Illinois Bufo amer- cantis americanus. Figures in parentheses are numbers of specimens. Characteristic Extreme Northern Illinois (11) Rans;e nout-vent length (mm.) lead length as per- centage ot body length 'arotoid length as percentage of body length 'entral pattern*. . . . )ne wart per dorsal spot 55.2-76,2 26.8-30.9 17.5-22.! 3-5 Mean 28.6 19.4 3.9 Per Cent of Specimens 64 Champaign- Sangamon Counties (17) Range Mean 60.8-90.0 26.7-30.2 15.6-21.5 3-5 28.6 19.0 4.1 Per Cent of Specimens Coles County (12) Range 55.2-74.8 26.4-32.9 16.6-23.4 2-5 Mean 29.2 20.4 3.0 Per Cent of Specimens 35 58 Adams- Hancock Counties (7j Range Mean 47.0-67.0 27.6-31.2 18.8-23.8 2-4 29.9 21.4 3.2 Per Cent of Specimens 100 *Ventral pattern indicated by index number. The index numbers have been assigned as follows: venter immaculate r with a single pectoral spot, I ; a few scattered spots on the breast region, 2 ; a zone of dark spots across the breast, ; entire venter spotted, 4; venter predominantly dark, 5. s at night. The female lays several thou- and eggs in long strings, and these strands nay extend for several feet along the bottom f a pond or ditch. Each jelly string cnm- lonly contains double or sometimes triple ows of eggs, the individual eggs ranging rom 1.2 to 1.4 mm. in diameter. Hatching iccurs in approximately a week. The small ilack tadpoles, easily recognized as toad arvae, fig. 53, are almost identical in all he Illinois species of Bufo. Metamorphosis ccurs in early June. Heavy rain late in the season may stimu- ate singing by small, newly matured males. Illinois Distribution.—With the excep- lon of the sand areas, where Bufo wood- \ousei fowleri predominates, B. a. ameri- flnus is the common toad of the northern lalf of Illinois, fig. 62. In much of the black oil prairie it is the only toad. Although his form is less conspicuous in northern lUi- ois than fowleri appears to be in the south- B. a. americanus, since both breeding dates and locality preclude the possibility of his specimens being foiuleri. Although undocumented by specimens, published records for the following locali- ties are believed valid ; most are indicated on the distribution map by hollow symbols: Cook County: Calumet City (Schmidt & Necker 1935) ; Evanston, Morton Grove (Necker 1939f ) ; West Pullman (Hubbs 1918); Wilmette (Necker 1939f ) ; Win- netka (Hubbs 1918); Wolf Lake (Schmidt & Necker 1935) ; Du Page County: Down- ers Grove, Glen EUyn, Naperville (Schmidt k Necker 1935); Kane County: Batavia, West Dundee (StiUe & Edgren 1948) ; Lake County: Fox Lake, Half Day (Schmidt & Necker 1935) ; Waukegan (Hubbs 1918); Stephenson County: Freeport (H. Garman 1892) ; Will Coun- ty: Joliet (Hubbs 1918); New Lenox (Schmidt & Necker 1935). 74 Illinois Natural History Survey Bulletin Vol. 28, Art. 1 Bufo americanus charlesmithi Bragg Dwarf American Toad Bufo terrestris charlesmithi Bragg 1954:247 (type locality: 1.8 mi. S, 7 mi. E Norman, Cleveland County, Oklahoma). Bufo lentiginosus americanus nee Holbrook, PYarrow 1882^:166 (part). fBufo americanus nee Holbrook, Burt & Burt 1929:2. Bufo americanus americanus nee Holbrook, Cagle 1942a:179. Diagnosis.—A small to medium-sized race of Bufo americanus (largest Illinois specimen 69 mm. from snout to vent but usually less than 60 mm. at sexual matur- ity), fig. 61, differing from B. a. ameri- canus in the following characteristics: smaller size; plain or sparsely spotted ven- ter; head length usually more than 30 per cent of snout-vent length (in 80 per cent of specimens) ; somewhat greater parotoid length; dorsal spots, if present, small and including a single wart (83 per cent of speci- mens). Remarks.—Although the dwarf form (B. a. c!iarles?riithi) and the large form {B. a. americanus) are difficult to distinguish because of the overlap in characters, there is little doubt that these two forms repre- sent distinct biological entities in Illinois. Whatever its status, the dwarf race has a considerable range. Parker (1939) illus- trated a specimen from Reelfoot Lake, Ten- nessee, where it is reportedly uncommon, and I have seen specimens in central Arkan- sas and southern Missouri. Variation.—Like the large form, B. a, charlesmithi is characterized by sexual di- morphism. Also, there is some evidence that in B. a. charlesmithi the males tend to be unicolorous, whereas females have obscure, dorsolateral light bands on a reddish ground- color. No information is available on ontogenetic variation. The few juvenile specimens avail- able are small replicas of the adults. Geographic variation for three samples is summarized in table 17. The number, length, and proportions are for sexually ma- ture specimens. The variation in dorsal and ventral patterns, however, is for both adults and juveniles. Habits.—The dwarf American toad, un- common in Illinois, is apparently a forest animal. Otherwise, it is probably similar to the large form in habits. Breeding aggre- gations of the small form have been encoun- tered on a few occasions in flooded fields and shallow ditches on the Mississippi River floodplain. Most specimens at hand, how- ever, have been found in steep-sided excava- tions that were too deep for the toads to climb or jump from. The ecological rela- tionships of this toad with Fowler's toad need study. The peak of the breeding season appears to be the first half of April. The call is a prolonged, high trill that I cannot distin- guish from the song of the large form. The eggs are laid in long strings, and there is a single row of eggs within each string. The individual eggs range from 1.5 to 2.0 mm, in diameter. Additional observations are Table 17.—Geographic variation in Illinois Bufo americanus charlesmithi. Figures in parentheses are numbers of specimens. Characteristic November, 1961 Smith : Amphibians and Reptiles of Illinois 75 needed to confirm the apparent difference between the two forms in number of rows of eggs per strand. I have not seen tad- poles, but it is likely that transformation occurs in late May and early June. Illinois Distribution.—The occurrence of B. a. charlesmithi is sporadic, fig. 62 ; this toad is subordinate in numbers to B. woodhousei fowleri even in the forested Shawnee Hills, where it is best known. Since adequate samples of B. americanus are not available from the Southern Division, the subspecific status of the toads from this area is uncertain. The rarity and the con- sistently small size of the few specimens known from this part of the state strongly suggest the dwarf race. Fig. 62.—Distribution of Bufo atncriccuius. Vertical hatching indicates the presumed range of Bufo americanus americanus in Illinois; horizontal hatching, the presumed range of Bufo americanus charlesmithi ; crosshatching, the area of intergradation between the two subspecies; solid circles indicate localities rep- resented by specimens examined during this study; open circles, localities represented by published records believed to be valid. The lower map depicts the total range of the species in the United States. The Richland County records of Cope (1889) and Gaige (1914) have not been accepted here, because Cope's Olney mate- rial that is still extant contains only B. iv. foivleri. Moreover, Mrs. Gaige in her paper failed to mention Fowler's toad, which is so abundant in Richland County that it is in- conceivable it would have been overlooked. There is some doubt concerning the Marion County record of H. Garman (1892) and the Washington County record of Burt & Burt (1929), but since there are recent col- lections of B. americanus in Bond, Clinton, Macoupin, and Montgomery counties. Gar- man's and Burts' records may be valid. No records for either race of americanus are known for the Wabash Border division. Although undocumented by specimens, published records for the following locali- ties are believed valid and are indicated on the distribution map by hollow symbols: Madison County: (Hurter 1911); Ma- rion County: Centralia (H. Garman 1892) ; St. Clair County: (Hurter 1911) ; Washington County: Nashville (Burt & Burt 1929). Bufo woodhousei fowleri Hinckley Fowler's Toad Bufo fowleri Hinckley 1882:310 (type locality: Milton, Massachusetts) ; Hubbs 1918:42-3. Bufo ivoodhoiisii foivleri, Necker 1938:2. Bufo woodhousei fowleri, Schmidt 1953:67. Bufo Icntiginosus americanus nee Holbrook, PYarrow 1882