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Autecology of the Copperhead BY O 1^ .1960 University of Kansas Publications „,,,.. , Museum of Natural History Volume 13, No. 4, pp. 85-288, pis. 13-20, 26 figs, in text November 30, 1960 Autecology of the Copperhead BY HENRY S. FITCH University of Kansas Lawrence 1960 UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY Institutional libraries interested in publications exchange may obtain this series by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas. Copies for individuals, persons working in a particular field of study, may be obtained by addressing instead the Museum of Natural History, University of Kansas, Lawrence, Kansas. There is no provision for sale of this series by the University Library, which meets institutional requests, or by the Museum of Natural History, which meets the requests of individuals. However, when individuals request copies from the Museum, 25 cents should be included, for each separate number that is 100 pages or more in length, for the purpose of defraying the costs of wrapping and mailing. * An asterisk designates those numbers of which the Museum's supply (not the Library's supply) is exhausted. Numbers published to date, in this series, are as follows: Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950. *Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 figmes in text. April 9, 1948. Vol. 3. *1. The avifauna of Micronesia, its origin, evolution, and distribution. By Rol- lin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951. *2. A quantitative study of the nocturnal migration of birds. By George H. Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951. 3. Phylogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 473- 530, 49 figures in text, 13 tables. October 10, 1951. 4. Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr., and Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10, 1951. Index. Pp. 651-681. *Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31 figures in text. December 27, 1951. Vol. 5. Nos. 1-37 and index. Pp. 1-676, 1951-1953. *Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. By Stephen D. Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10. 1952. Vol. 7. *1. Mammals of Kansas. By E. Lendell Cocknmi. Pp. 1-303, 73 figures in text, 37 tables. August 25, 1952. 2. Ecology of the opossum on a natural area in northeastern Kansas. By Henry S. Fitch and Lewis L. Sandidge. Pp. 305-338, 5 figiu-es in text. August 24. 1953. 3. The silky pocket mice (Perognathus flavus) of Mexico. By Rollin H. Baker. Pp. 339-347, 1 figure in text. February 15, 1954. 4. North American jumping mice (Genus Zapus). By Phillip H. Krutzsch. Pp. 349-472, 47 figiu-es in text, 4 tables. April 21, 1954. 5. Mammals from Southeastern Alaska. By Rollin H. Baker and James S. Findley. Pp. 473-477. April 21, 1954. 6. Distribution of Some Nebraskan Mammals. By J. Knox Jones, Jr. Pp. 479- 487. AprU 21, 1954. 7. Subspeciation in the montane meadow mouse, Microtus montanus, in Wyo- ming and Colorado. By Sydney Anderson. Pp. 489-506, 2 figures in text. July 23, 1954. 8. A new subspecies of bat (Myotis velifer) from southeastern California and Arizona. By Terry A. Vaughan. Pp. 507-512. July 23, 1954. 9. Mammals of the San Gabriel mountains of California. By Terry A. Vaughan. Pp. 513-582, 1 figure in text, 12 tables. November 15, 1954. 10. A new bat ( Genus Pipistrellus ) from northeastern Mexico. By Rollin H. Baker. Pp. 583-586. November 15, 1954. 11. A new subspecies of pocket mouse from Kansas. By E. Raymond Hall. Pp. 587-590. November 15, 1954. 12. Geographic variation in the pocket gopher, Cratogeomys castanops, in Coa- huila, Mexico. By Robert J. Russell and Rollin H. Baker. Pp. 591-608. March 15, 1955. 13. A new cottontail (Sylvilagus floridanus) from northeastern Mexico. By Rollin H. Baker. Pp. 609-612. AprU 8, 1955. 14. Taxonomy and distribution of some American shrews. By James S. Findley. Pp. 613-618. June 10, 1955. 15. The pigmy woodrat, Neotoma goldmani, its distribution and systematic posi- tion. By Dennis G. Rainey and RoUin H. Baker. Pp. 619-624, 2 figures in text. June 10, 1955. Index. Pp. 625-651. (Continued on inside of back cover) University of Kansas Publications Museum of Natural History Volume 13, No. 4, pp. 85-288, pis. 13-20, 26 figs, in text November 30, 1960 Autecology of the Copperhead BY HENRY S. FITCH University of Kansas Lawrence 1960 University of Kansas Publications, Museum of Natural Histort Editors: E, Raymond Hall, Chairman, Henry S. Fitch, Robert W. Wilson in test Volume 13, No. 4, pp. 85-288, pis. 13-20, 26 Bgs. Published November 30, 1960 Mils. COlVir. iLyOl t DEC 2 7 1960 . University of Kansas Lawrence, Kansas PRINTED IN THE STATE PRINTING PLANT TOPEKA. KANSAS 1960 28-4428 Autecology of the Copperhead BY HENRY S. FITCH CONTENTS PAGE Introduction 89 Acknowledgments 92 Methods 93 Description Lepidosis 99 Color and Pattern 102 Size 103 Bodily Proportions 106 Dentition 108 Hemipenis 112 Relationships 113 Habitat 116 Range and Geographic Variation 121 Behavior Crawling 124 Coiling 126 Swimming 128 Climbing 128 Disposition 129 Combat Dance 131 Shedding 134 Hibernation and the Effect of Temperature 137 Movements 147 Reproduction Courtship and Mating 157 Fecundity of Females 162 Development of Ova and Embryos 164 Aggregating of Gravid Females 166 Time of Birth 168 Number of Young per Litter 171 Birth of Young 176 Behavior of Females 17S Defects and MortaUty at Birth 178 The Egg Tooth 179 (87) 88 University of Kansas Publs., Mus. Nat. Hist. FACE Size at Birth 181 Appearance of Young 182 Growth and Development Utilization of Stored Yolk, and Early Growth 183 Later Growtli 183 Cessation of Growth 192 Food Habits Methods of Obtaining Prey 193 Luring of Prey by Young 196 Statements of Food Preferences 198 Composition of the Diet 199 Kinds of Prey 201 Amount of Food Consumed 211 Defense, Escape and Mortality Factors Defense and Escape 219 Natural Enemies and Predation 221 EflFects of CHmatic Extremes 226 Parasites, Diseases and Injuries 228 Composition of the Population 230 Numbers 236 Relation to Man Attitudes of tlie Pubhc 245 Survival Under Modem Conditions 249 Control 250 The Venom and Bite Adaptations Correlated with the Venom 253 Properties of the Venom 255 Quantity of Venom Produced 256 Toxicity of tlie Venom 256 Susceptibilit)' of Snakes 259 Circumstances and Outcomes of Bites 261 Hypersensitivity to Venom 262 Case History of a Bite 263 Treatment of the Bite 265 Summary 269 Literature Cited 277 INTRODUCTION In 1948 when ecological studies were initiated on tlie newly created University of Kansas Natural History Reservation, the copperhead was one of the first species that attracted attention as meriting intensive investigation. As an abundant predator on small vertebrates, including both those that are primary consumers of the vegetation and those of higher trophic levels, it was recognized as a key animal in the local ecosystem. Despite persistent effort to study the copperhead, progress was slow, especially in the early stages of the investigation. Copper- heads were rarely seen engaged in their normal activities, and even when such individuals were found, observing them proved to be remarkably unrewarding. A copperhead found by chance usually lay motionless for long periods, either having "frozen" in the usual reaction to any alarm, or merely resting—the sluggish behavior that is characteristic of the species. Attempting to ob- serve such a snake severely tried the patience of the investigator. When the snake finally began to move, it might soon be irretrievably lost because of the perfection with which it blended with its back- ground, and the dense concealing vegetation and other cover in the situations frequented. In the summer of 1949 cylindrical wire funnel traps set for lizards at the edges of rock outcrops in the "Rat Ledge" area caught several copperheads, and many more of these snakes were trapped in similar situations in autumn of the same year. Thereafter, each autumn, trap lines were maintained in rock ledge habitat and cop- perheads were obtained in numbers at that time of year but not at other seasons. In 1957 trap lines were established in a variety of habitats not previously sampled, and tliis trapping was continued on a larger scale in 1958 and 1959. In tliese three years copper- heads were obtained in large numbers tluroughout the season of their activity. The present report is based primarily upon records obtained on the 590-acre University of Kansas Natural History Reservation, in the nortlieas tern corner of Douglas County, Kansas, and on the adjacent 160-acre Rockefeller Experimental Tract. Including a few miscellaneous records, such as tliose of snakes found dead on county roads, and of young bom dead in captivity, a total of 1532 (89) 90 University of Kansas Publs., Mus. Nat. Hist. individual copperheads obtained from the Reservation or imme- diately adjacent areas were recorded a total of 2018 times between July 1, 1948, and November 9, 1959. Supplemental information was obtained from numerous other copperheads collected or ob- served elsewhere in eastern Kansas, notably from large series taken near La Cygne by Vernon Mann, who kindly permitted me to examine the live snakes in his possession from time to time. Despite the rapid accumulation of data during the later years completion of the study was long delayed because of seeming in- adequacies or inconsistencies in the information obtained.
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