auinxd OF THE Bee Best mVGkyaeliaies PAZOUROR Ie. 0e Sura ese Jonathan A. Campbell-and William W.-LamarTHE VENOMOUS REPTILES ‘OF THE VOLUME II WESTERN HEMISPHERE JONATHAN A. CAMPBELL University of Texes at Arlington WILLIAM W. LAMAR University of Texas af Tyler with contributions by EDMUND D. BRODIE Ill and EDMUND D. BRODIE JR. Indiana University and Uta State University RONALD L. GUTBERLET JR. and MICHAEL B. HARVEY LUnizesty of Texas at Tyler and East Tess State University ROBERT NORRIS Stanford Univesity Medical Centr DAVID A. WARRELL, Conte for pice Medicine, Unioesity of Oxf VINICIUS XAVIER DA SILVA Universidade de So Paulo Comstock Publishing Associates 4 pivision or conwe1t. UNIVERSITY PRESS, ITHACA AND LONDONCopyright © 2004 by Comell University All rights reserved, Except for bref quotations ina review his bool, oF parts theveof, mast not be repreuced in any form without permission in writing from the publisher. For information, address Cornell Univesity Press, Sage House, 512 East State Stroct, Ithaca, New York 14850 First published 2004 by Comell Unversity Press Print in China Libary of Congres Catalging-in-Publiation Data Campbell, Jonathan A ‘The venomous repiles ofthe Wester Hemisphere / by Jonathan Campbell and Wiliam W. Lamar, with contributions by Edmund D. Brodie I... et al ._ cm —{Comstock books in herpetology) Inchides bibliographical references and index ISBN 0-8014-41412 (Cloth :alk. paper) 1, Poisonous snakes—Wester Hemisphere. 2. Reptiles—Western Hemisphere. 1 Lamar, William W, 1950- IL Brodie, Edmund D, 1963- ML Tile. IV Series. OLss6.06C24 2008 587 9616509181221 20200783 Cornell Unversity Press strives to use environmentally responsible suppliers and materials to the llet extent possible inthe publishing ofits books, Such materials inchide vegetable-based,lows-VOC inks td acide papers that ae recycle, totally chlorino-ee, or partly compos of nomwood fibers. For father information, vist our ‘website a wws.comellpress comell Cloth priming 10987654321We dedicate this book to our wives, Tanya Dowdey and Nancy Lamar, who have enthusiastically supported our research activities.CONTENTS VOLUMEIL Introduction 1 REGIONAL ACCOUNTS AND KEYS TO VENOMOUS REPTILES Canada and the United States u Canada, 11. United States, 12 Mexico and Central America 18 Mexico, 19. Guatemala, 34. Belize, 36, Honduras, 38. El Salvador, 40. Nicaragua, 41. Costa Rica, 48, Panama, 47. ‘The Caribbean Islands 49 ‘The Windward Islands, 49. The Netherlands Antilles, 50. Islas Los Testigos, 50. Trinidad, 50. South America 51 Colombia, 51. Venezuela, 58. Guyana, 61. Suriname, 62. French Guiana, 64. Brazil, 65. Ecuador, 75. Peru, 78. Bolivia, 34. Paraguay, 86. Uruguay, 88. Argentina, 89. TAXON ACCOUNTS i Venomous Lizards, Family Helodermatidae 95 Gila Monster and Beaded Lizard, Genus Heloderma Wiegmann, 1829 95 Heloternw horridum (Wiegmann), 103. Heloderma suspectum Cope, 106. Coralsnakes and Seasnakes, Family Elapidae 108 Coralsnakes 108 Slender Coralsnakes, Genus Leptomicrurus Schmidt, 1937 125 Leptomicrurus collaris (Schlegel), 128. Leptomicrurus navduecti Jan), 129. Leptonsicrurus renjifo Lamar, 130. Leplomticrurus seutiventris (Cope), 130. Sonoran Coralsnake, Genus Micruroides Schmidt, 1928 132 Micruraides euryxanthus (Kennicott), 133 Coralsnakes, Genus Micrurus Wagler, 1824 135 The Monadal Group. 45 Micrurus albicinctus Amaral, 145. Micrurus alleni Schmidt, 146. Micrurus anvellatus (Peters), 148. Micrurus averyi Schmidt, 149. Micrurus bernadi (Cope), 150. Micrurus bocourti Jan), 151. Micrurus bogerti Roze, 152“Micrurus brown Schmit and Smith, 153. Micrarus catamayensis Roze, 155. Micruruscircinais (Duméri and Bibron), 156. Micrurus claxki Schmit, 158. Micruvus corallinus (Merrem), 159. Micrurusdiastema {(Damért, Bibron, and Duméril), 162. Micrurs dstans Kernicot, 163. Micrurus dumeri Qan), 16 Micruras epkippfer (Cope), 168, Micrarus fuloius (Linnaeus), 169. Micrurushippocrepis (Peters), 171 Micrurustnigedorfi Wagler, 172. Micriruslatiascats Schmidt, 174. Micrurus linbatus Frasen, 174 Micrurus margartiforus Roze, 175. Micrurus medemi Roze, 175. Micrarus mertensi Schonidt, 177. Micrurus hebulans Rose, 177. Micrurux nigrocinctus (Girard), 179. Micrurs oligonnelitus Ayerbe and Lope, 183 ‘Micrunus ornatissinis Jan), 183. Micrurus pachecogili Campbell, 184. Micrurus paraensis Cunha and Neccmento, 188, Micrurus peruoianus Schmidt, 286. Micrurus petersi Roze, 187. Micrurus proximarts Smith and Cheapliry, 187. Micnous psyches (Daudi), 189. Micrurs patumeyensis Lancins, 190, Mvurus reals Roze, 100) Micrurus ruatanus (Giinther), 192. Micrurus sanglensis Nicéforo-Maria, 191. Mieruras Sieindachneri (Wenner), 192. Micrurus stewart Barbour and| Amaral, 193. Micrutus stuarti Roze, 19. Micryyus tener (Baird and Girard), 195. ‘The Central American Triad-Bearing Group 197 Micrurus elegans (an), 197. Micrurus laticollaris (Peters), 199. ‘The Bicolored Group 199 Micrurs nipartitus (Duméil, Bibron, andl Duméril), 200. Micrurus mulifescitus jan), 202. Micrurus ‘multiseutatus Rendahl and Vestergren, 204. Micrurus spurrelti(Boulenger), 204 ‘The South American Triad-Bearing Group 205 “Micrurusaltirostris (Cope), 205. Micrurus ancoralis (an), 208.’ Mierurus baliocoryphus (Cope), 209. Micrurus trasliensis Roze, 210. Micrurus decoratus (an), 211. Micrurus diana Reze, 212, Micrurus dissoleucus (Cope), 213. Micrurusfiiformis (Giinther), 215. Micrurus frontalis (Duméril, Bibron, and Duméril), 217. Micrurus hemprcki Jan), 217. Micrurus iboboca (Merrem), 219. Micrurus isozous (Cope), 220. Mieruras ienisalus (Linnaeus), 221. Micrurus meridensis Roze, 224. Micrurus pyrttocryplus (Cope), 225. Micrurs serrams Harvey, Aparicio-E, and Gonzélez-A., 226, Micrurusspsii Wagler, 227. Micrurus surinamensis (Cuvier) 23). Micrurus tschualt (Jan), 232. Seasnakes 233 Yellow-Bellied Seasnake, Genus Pelamis Daudin, 1803 233 Pelamis platurus (Linnaeus), 236 Pitvipers, Family Viperidae 240 Contents Cantils, Copperhead, and Cottonmouth, Genus Agkistrodon Palisot de Beauvois, 1799 249 Aghisradonbitinerts Conther, 262. Agisiradon taylor Bunger andl Robertson, 265. Agkistoon contortis (Linnaeus), 266. Agkistradon piscicorus (Lacépede), 271 Jumping Pitvipers, Genus Atropoides Werman, 1992 274 “Atropides mexicenus (Duméril, Bibeon, ane! Duméri, 281, Alrpoides numifer (Rappell) 284. Alropoides sceiduns (Hoge), 285. Atropoides olmee (Pérer-Higareda, Smith, and Julié-Zertuche), 27. Atropides pcadoi (Dunn), 288, Palm-Pitvipers, Genus Bothriechis Peters, 1859 290 Bothriechis aurfer (Salvin), 296. Bothreckis bicolor (Bocourt), 297. Bothricis laterals Peter, 299. Bothricis nari (Barbour and Loveridge), 30L. Bothrieohis nigrovrids Peters, 302. Bothriechis rowley (Bogert), 308. Bothrechis shlegli (Beethold), 304, Bothriechis supraclinrs (Taylor), 307. Bothrechis Uulasinus Campbell and Smith, 308. Forest-Pitvipers, Genus Bothriopsis Peters, 1861 309 uthropaishilienta(Wied-Newssied, 313. Bothrops coromelas(Boulenger), 315. Bothrops mesa {Sternfeld), 376. Bothriopsisoligoepis (Werner) 318. Bothriopsis pulchra (Peters), 319. Bothriopsis tneniata (Wagler), 320 Toadheaded Pitvipers, Genus Bothrocophias Gutberlet and Campbell, 2001 322 Bovhrocophias compel (Freire-Lascano), 326. Bothrocophiascolombianus (Rendahl ancl Vestergren), 329. Bothrocophins hyoprra (Amaral), 380. Botlyocophias micrphthatrnes (Cope), 332. Bothrocophins meversh Gutberlet and Campbell, 33.Lanceheads, Genus Bothrops Wagler, 1824 3a Bothrops aleatraz Marques, Martins, and Sazima, 364, Bothrops alternatus Duméri, Bibron, and Duméril, 365. Bothrops ammadytoes Leybold, 368. Bothrops andianus Amaral, 369. Bothrops asper (Garman), 371 Bothrops atrox (Linnaeus), 376, Bobs barnetti Parker, 380. Bothrops brazili Hoge, 381. Bothrops caibbaeus (Garman), 383. Bothrops cotiara (Gomes), 385. Bothrops erythromclas Amaral, 386. Bothrops fonseeai Hoge and Belluomini, 388. Bothrops insularis (Amaral), 388, Bothrops itaetinbygae (Boulenger), 38%. Bothrops jararaca (Wied-Neuwied), 390. Bothrops jararacussu Lacerda, 392. Bothrops jonathani Harvey, 3%. Bothrops Janceolatus (Bormaterte), 395. Bothrops leucurus Wagler, 397. Bothrops lojantus Parker, 399. Bothrops tmarajoensis Hoge, 400. Bothrops moojeni Hoge, 400. Bothrops muriciensis Ferrarezzi and Freire, 401. Bothrops oshornei Freire-Lascano, 402. Bothrops pictus (Tschudi), 404, Bothrops pirajai Amaral, 405. Bothrops punctatus (Garcia), 406. Bothrops roedingeri Mertens, 407. Bothrops sanctaecrucis Hoge, 408. Bothrops venezuelensis Sandner Montilla, 409. ‘The Bothrops newwied Complex by Vinicius Xavier da Silva 410 Bothrops neusviedi Wagler, 11. Botlrops diporus Cope, 413. Bothrops lutzi (Miranda-Ribeiro), #15 Bothrops mattogrossensis Amaral, 417. Bothrops pauloensis Amaral, 418. Bothrops pubescens (Cope), 419 Bothrops sp. 421. Montane Pitvipers, Genus Cerrophidion Campbell and Lamar, 1992 422 Cerroptiion tarbouri (Dunn), 429. Cerrophidion godmani (Giinther), 431. Cerraphidion petlatcalensis Léper- Luna, Vogt, and Torre-Loranca, 434. Cerrophidion tzotzilorum (Campbell) 435. Bushmasters, Genus Lackesis Daudin, 1803 436 Lachesis acrochorda (Garcia), #42. Lachesis melenocephala Solbrzano and Cerdas, +. Lachesis muta (Linnaeus), 445. Lachesis stenophrys Cope, 448. Mexican Horned Pitvipers, Genus Oplryacs Cope, 1887 449 Opliryacus melanurus (Miller), 452. Ophryacs undulatus (Jan), 454. Hognosed Pitvipers, Genus Porthidium Cope, 1871 456 Posthidian arcosae Schiitti and Kramer, 461. Portidivan dunn (Hartweg and Oliver), 462, Porthidium iespere (Campbell), 464, Porthidium lonsbergii (Schlegel), 465. Portidium nasutum (Bocourt), 467. Posthidium ophryomegas (Bocourt) 470. Porthidium porrasi Lamar and Sasa, 472. Porthidiwn voleanicum Solérzano, 475. Porthidium yuentanicim (Smith), 474. Index (1) VOLUME II Rattlesnakes a7 Rattlesnakes, Genus Crotalus Linnaeus, 1758 490 Crotalus adamiantews Palisot de Beauvois, 525. Crotalus aquilus Klauber, 527. Crotalus atvox Baird and Girard, 528, Crotalus bnsiliscus (Copel, 543. Crofalus catalinensis Cliff, 536, Crotalus cerastes Hallowell, 537. Crotalus durissus Linnaeus, 539, Crotalus enyo (Cope), 547. Crotalus horridus Linnaeus, 548. Crotalus intermedius Troschel, 552. Crotalus lannmi Tanner, 554. Crotalus lepidus (Kennicott), 995. Crotalus mitchell (Cope), 558. Crotalus molossus Baird and Gieard, 561. Crotalus oreganus Holbrook, 564. Crotalus polystictus (Cope), 570. Crotalus pricei Van Denburgh, 572. Crotalus pusilius Klauber, 574. Crotalus rows Cope, 57. Crotalus ruber Cope, 377. Crotalus scutulatus (Kennicolt), 579. Crotalus simus Latrelle, 583. Cro‘alus stejnegeri Dunn, 586. Crotalus figris Kennicott, 587. Crotalus tortugensis Van Denburgh and Slevin, 589, Crotalus totonecus Gloyd and Kautfeld, 590. Crotalus transversus Taylor, 591. Crotalus triseritus (Wagler), 592. Crotalus viridis (Rafinesque), 594. Crotalus willardi Meek, 597. Massasauga and Pygmy Rattlesnake, Genus Sistrirus Garman, 1884 oor Sistrurus catenatus (Rafinesque), 609. Sistrurus milavius (Linnaeus), 614 Contents ixContents Venomous Snake Mimicry 617 by Edmund D. Brodie III and Edmund D. Brodie fr. ‘The Evolution of New World Venomous Snakes 634 by Ronald L. Gutberlet Jr. and Michael B. Harvey Venom Poisoning by North American Reptiles 683 bby Robert Norris Snakebites in Central and South America: Epidemiology, Clinical Features, and Clinical Management 7019 by David A. Warrell Glossary 763 Literature Cited 75 Index {1} Color Maps are betiveen pages 44 and 45 (Vol. 1) Color Plates 1-751 are between pages 92 and 93 (Vol. 1); 752-1365, pages xvi and 477 (Val. ID; 1366-1500, pages 684 and 685 (Vol. ID)TABLES BEBRESBENRREREE VOLUME 1 Distributions by physiographic division or provinee of venomous reptiles in Canada and the United States 12 Distributions by vegetational association of venomous reptiles in Canada and the United States 13 Distributions by physiographic region of venomous reptiles in Mexico 22 Distributions by vegetation type of venomous reptiles in Mexico. 24 Distributions by vegetation type of venomous reptiles in Guatemala 35 Distributions by district of venomous snakes in Belize 37 Distributions by vegetation type of venomous snakes in Honduras 39 Distributions by vegetation type of venomous snakes in Fl Salvador 41 Distributions by vegetation type of venomous snakes in Nicaragua 42 Distributions by vegetation type of venomous snakes in Costa Rica 45 Distributions by vegetation type of venomous snakes in Panama 48 Distributions by vegetation type of venomous snakes in Colombia 53 Distributions by vegetation type of venomous snakes in Venezuela 59 Distributions by vegetation type of venomous snakes in Guyana 61 Distributions by vegetation type of venomous snakes in Suriname 63 Distributions by vegetation type of venomous snakes in French Guiana 64 Distributions by physiographic magion of venomous snakes in Brazil 67 Distributions by vegetation type of venomous snakes in Brazil 68 Distributions by vegetation type of venomous snakes in Ecuador 76 Distributions by physiographic region of venomous snakes in Pert 0 Distributions by physiographic region of venomous snakes in Bolivia 85 Distributions by physfographic region of venomous snakes in Paraguay 87 Distributions by physiographic region of venomous snakes in Uruguay 88 Disiributions by physiographic region of venomous snakes in Argentina 90) Variation in selected scale characteristics in species of Heloderma 98 ‘Variation in selected features in species of Leplomicrurus 110 Variation in selected features in species of Micrurus 111 Comparison of certain morphological features of New World genera of pitvipers 242 Variation in selected scales in species of Agkistradon 250 Variation in selected features in species of Atropoites 275 Variation in selected scale characteristics in species of Bothriechis 291 Measurements and dates of bitth for litters of captive-born Botliriecis schlegeli 25 ‘Variation in selected scale characteristics n species of Bothriepsis 310 Reproduction in Bothriopsistaeniata from Tepoe, Sipaliwini, Suriname 312 . Variation in selected features in species of olirocophias 32336, Variation in selected features in species of Bothrops 335 37, Measurements and dates of birth for captive-born young of Botivops allernatus produced by a single female 351 58, Reproduction in captive Bothrops neue (sensu lato) 354 59, Variation in selected scale characteristics in species of Cerrophidion 423 40, Number and frequency of prey items recovered from Cerrophidion gadmani from Nuclear Central ‘America $25 41, Size and variation in litters of Cerrophidion godmani from Guatemala and Oaxaca, Mexico 427 42, Variation in selected characteristics in species of Lachesis 438 4, Variation in selected scale characteristics in species of Ophryacus 450 “44, Variation in selected scales in species of Porthiiwm 457 VOLUME I 45. Variation in selected cephalic scale characteristics in species of Crotalus 492 46, Variation in certain characteristies in species of Crotalus 495 47, Number of Crotalus horridus bountied in selected years in Houston County, Minnesota 504 48. Prey recovered from Crotalus atrox in Texas 508 49, Reproduction in Crotalus 517 50. Comparison of selected features of the Crotalus durissus group 546 SL. Variation in subspecies of Crotalus lepidus 557 Variation in selected scale characteristics in rattlesnakes of the genus Sistrurus 602 Genera of venomous snakes of the Americas and their potential mimics 625 54. Allocation of Bothvops (sensu lato) species to five genera by Burger (1971) 656 55, Revision of Porthidivn from 1971 to the present 657 56. Allocation of Crotalus species to seven groups by Murphy ot al. (2002) 667 57. Venom yields of some North American venomous reptiles 65 58, Clinical grading scale for pitviper bites 700 59, Starting doses of antivenoms commercially available in the United States 702 60. Distribution of rear-fanged or mildly venomous colubrid snakes by genus in North and Central America, including islands 719 Distribution of rear-fanged or mildly venomous colubrid snakes by genus in the Caribbean region 719 (62. Distribution of rearfonged or mildly venomous colubrid snakes by genus in South America, including, islands 720 63, Antivenoms available for treatment of envenoming by American snakes 756 TablesABBREVIATIONS FHGO FMNH FUL. HINIRENA 1B INKMT. 1B JAC JMR ; ; ; AMNIT ANSP ace BMNH BYU cas cor crRaD co cr sa cvuce cvuLa DMNH ENEPL ENS FeD. | MENG MHNUC MNHN MNRI American Museum of Natural History, New York, New York, USA. Academy of Natural Sciences of Philadelphia, Philadelphia, Pennsylvania, USA Bryce C, Brown collection (now Strecker Museum), Waco, Texas, USA British Museum of Natural History, London, United Kingdom Brigham Young University, Provo, Utah, USA California Academy of Sciences, San Francisco, California, USA Coleccién Boliviana de Fauna, La Paz, Bolivia Centre de Coopération Internationale en Recherche Agronomique pour le Développement, French Guiana Camegie Museum, Pittsburgh, Pennsylvania, USA Colegio Paralela of the Instituto Butantan, So Paulo, Brazil Coleccién Santiago Ayerbe Colegao da Vertebrados, Universidade Catélica de Goi, Brazil Coleccién Vertebratos, Universidad de los Andes, Mérida, Venezuela Dallas Museum of Natural History, Dallas, Texas, USA Escuela Nacional de Estudios Profesionales Iztacala de la Universidad Nacional Autsinoma de México, Mexico Erie N, Smith collection Fundagio Ezekial Diaz, Belo Horizonte, Fundacisn Herpetologica Gustavo Oreés, Quito, Feuador Field Museum of Natural History, Chicago, Ilinois, USA Federacion Universitaria de Litoral, Argentina Hexpetologfa, Instituto de Investigaciones sobre los Recursos Naturales, Universidad! Michoacana de San Nicolas de Hidalgo, Michoacan, Mexico o Paulo, Brazil 10 Paolo, Brazil Instituto Butantan, Instituto de Bigiene y Medicina Tropical, Guayaquil, Eeuador Instituto Vital Brazil, Rio de Janeiro, Brazil Jonathan A. Campbell collection Juan Manuel Renjifo collection Museum of Natural History, University of Kansas, Lawrence, Kansas, USA. Los Angeles County Museum of Natural History, Los Angeles, California, USA Laurie J. Vit collection Louisiana State University Muscurn of Zoology, Baton Rouge, Louisiana, USA Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA ‘Muséum d'Histoire Naturelle, Geneva, Switzerland Museo de Historia Natural, Universidadl de Cauea, Cauca, Colombia Muséum National d'Histoire Naturelle, Paris, France ‘Museu Nacional, Rio de Janeiro, BrazilMSNM MUFAL MvZ MZUSP NCMNH NHRM NMBA NMW NoRMAT PNN Qcaz RMNH RWM SDSNH STRI TeWC uce UCR PMT uUMMZ UNAMLT UNAM-NL UNESP usNM UTA uw WIN ww ZEMK iu ZMB ZMUU ZsBs ZsM ZUEC For journal abbreviations, see Abbreviations Museo Civico de Storia Naturale de Milano, Milan, Ttaly Museu Universidade Federal de Alagoas, Alagoas, Brazil Museum of Vertebrate Zoology, University of California at Berkeley, Berkeley, California, USA. Museu de Zoologia, Universidade de Sa0 Paulo, Si0 Paulo, Brazil [North Carolina Museum of Natural History, Raleigh, North Carolina, USA [Naturhistoriska Riksmuseet, Stockholm, Sweden [Naturhistorisches Museum, Basel, Switzerland [Naturhistorisches Museum, Vienna, Austria Niicieo de Offologia Regional de Mato Grosso, Parque Nacional Natural, Colombia Museo de Zoologia, Pontifica Universidad Catolica del Ecuador, Quito, Eeuador Natuuhistorisches Museum, Leiden, Netherlands, Roy W. MeDiarmid collection San Diego Society of Natural History, San Diego, California, USA ‘Smithsonian Tropical Research Institute, Panama “Texas Cooperative Wildlife Collection, Texas A&M University; College Station, Texas, USA Universidade Catélica de Gols, Goiss, Brazil Universidad de Costa Rica, San José, Costa Rica Universidad Federal de Mato Grosso, Mato Grosso, Brazil University of Michigan Museum of Zoology, Ann Arbor, Michigan, USA Universidad Nacional Auténoma de México, Los Tuxtlas, Veracruz, Mexico Universidad Nacional Autonoma de México, Nuevo Leén, Mexico Universidad Estadual Paulista, Brazil National Museum of Natural History, Washington, D.C, USA University of Texas at Arlington, Arlington, Texas, USA Universidad del Valle, Guatemala City, Guatemala Wilfred T. Neill collection William W. Lamar collection Zoologisches Forschungsinsti¢ut und Museum Alexander Koenig, Bonn, Germany Zoologisches Institut, Universitit Gottingen, Gottingen, Germany Universitit Hambolet, Zoologisches Museum, Berlin, Zoological Museum of the Royal University at Uppsala, Sweden Zoologische Sammlung des bayerischen Staates, Munich, Germany fermany Zoologische Staatssammlung, Munich, Germany Museu de Historia Natural, Universidade Estadual de Campinas, Campinas, St0 Paulo, Brazil erature Cited, For abbreviated terms, see Glossary.‘he Venomous Reptiles of the Western HemispherePlate 753. USA. Phos Photo by Toy D. om TL, UTA R Tafsir, Hi iy Dovid nama W. Porn Dade County, Flori, us, juvenile, 18 Sem TL, UTAR nt west of igus, Michoocdn,em TL, UTA R. Mexico, ‘elevation whe Plate 767. Crotalus aps, female, From 19km east of San Luis Potosi, Son Luis Potosi, Mexion, Photo by Dovid G Barker Plate 770. Collus ais, adult. Al Potosi Mexico, Photo by Louis W. Por Plate 773. C Rinconoda, Mi vt Thilem TL, ENS 10538. ila, Veracruz, Mexico, Photo by Eric N, Smith, courtesy of Robert Mora Galland Plate 765. Crofalas apis, TEWC west of Nich Guanajuato, Mexico, Ph 506, roe 33k by Robert A. 66. Cras apis, ‘Marmoles, 84 female, 42.0cm TL, UTA ithwest-of Dung, Hidalgo, Mexico, elvatio P ila, adit, El Lebo, Querétaro, “Mexico, Photo by Louis W. Poros Plate 769, Crotalus aus, adult, EL Valle de Fatasmas, San Ls Paton, Mexic, Photo by Lous} Pore 2i5mn.sino, juvenile. On Chicken Sprin rots alos, female. Appr othe junction with Alamo TL- Bend, Llano County, Tenas, US Asticus, mate, 19.0em TL, F ‘0 Phoio by Erie N. Manvel Varela: JulisPlate 788, Cries ctalnensic sla de Santa Ca 89, Croflusevllinenss Same as Plate 785, View Plate 790, Crotalus cetalinonss. sla de Santa Ct aj Califa Sur, Metco. Photo by Lous ftaane vestigial alle. Thisspecies and C. rubs forer- Uj Caliornia Sur, Meio. Photo by Paul Freed haracterstialy aed Ue ates, Photo by Louis Plate 791. Coteus rates, juvenile. On Plate 792. es artes. Southern Nevada, ms es croton RediosieRands Rond, km east of the junction with oto ima County, Anzona, USA. Phat Californian Highssoy 14 Keen County, Calfornio, USA. oto by Toy D. Hibbs te 795, Cra Lem TL RMNH Matapica Creck, comsal Suriname. Photo by Albertus Abu sit, Imperial (Count, California, USA. Photo by Louis W. Porras, Plate 797. Crt sacl. Brazil Photo by Plate 798 Ludwig Truman, ately 120, Ivan Savin Plate 798 Crotalus dais coliinsts, female, app imately 100em TL, IB, Catalso, Gods, Broil. PotPlate 807, z nen coustesy of I Mund de la Se milo Goel onsideably in Venezuuna, mle. Bow Vista, Plate B13. Crotalus 3 ion, Pate 814. Cr Roraio, Bret. fo Martins, Rosia, Broil. D ZZUEC S10. Co ‘omen, Campinas, So Fale, Bea Plate 815 ult female, LV 207 Alt Tail. Photo by Ture. Vit nicl: Aruba. Photo by Plate $20, Cra 1 ucolor, adlt fe Aruba. Photo by Lous W, Porras Irgonicns. Sipalwini Plate 819. Croat ‘surname. Photeby Joep Paul Free. dis, Las Sabanas to by Pal Pree Matarin, Manag sult female, Plate 822. Ci dis. Las Sabanas de re Dallas Zoo. Maturin, Mo o by Louis W Porn Aa, Photo byfe A Coihes s de Motrin, wis, neonate. Las Plate Venezuela, Photo by Mexico Hs enyo eyo, Baja California Norte, Plate 826. Cratlus enya emy, male, Mexico. Pho bby Ba Cassano, David. Barker Plate 828 yo fureus,Usja California Norte, Plate 829. Crofts 22358 aly Fis County, Texas, USA. Photo By David [Louis W. Portas at the Dall Zao, Plate 831 Cro Plate $32. Crt Carina, USA. Ph County, Noth C Devencer BES Crouluslrvidus, alt. Cherokee National a nn c termed interes. Nast of MoaiseCouny, Tennesse, USA. Photo by Terry ery Base, cour, male, Plate #40 Crialas fomoni, female Plate 84 ‘of Omiieme, men, 63am TL, BYU 2380 mately 45c 207m. Photo by jo Plate si tains Research Cent Photo by Toy D. Hibbs, wis, male. Indio Moun- Plate 83. Cras epi ul female. Vik Plate 814, Civil Val Verde Cov fspeth County, Texas, USA. of Butalo Trails Boy Seoul Camp on Ranch Rev Texas, USA. Photo by Louis W Porras. pits Kauteri, male, Chita, Keer, mae, appr 2h above, “Cas ha, Mesointans, Daa Ana County, New — Cathalral Visa Point Conyun, Chiicahua Mount ous W. Ports, (Cochise County, Arizona, UBA, el Am by Louis W. Peas Bhs ius Bieter, adult male. Rancho Pla c Aguila, Plate Beira, Darang, Mexico. Photo by Lowis W. Plate $8, ymale. From 17.1km Alazatas, Coahuila, Mexico, David G. Barker at the Plate 860, Cras eit tet ofS Antonia de Dills Zoo. Pate Tamaslip sulaul: Puesto Plate 867 H. Tashi Jo, Byja Calfvnia Norte, Me uns, adult, Rancho Vie, Niexco. Photo by Louis W. Use jn California Ni St the California Academy of Pate 562 alt feral ta Photo by Past Fe Pate 65. rath jh a tens, adult. I jenic. Photo by La Pate 8 nthe mo Plate 871. Cro rhae ical: pyres,subadul. From “ : adult. Furnace Plate Ta Rumarost Esjn California Norte, 3 northwest of the junction of Count Phi by Ty D. Hibbs Mesquite Valley Road and Smith Tale Road, Inyo adull. North- Plate $26, Cra slifrnia, US UTA RUNS Harguahala Mountains La Pa SA. Photo by Toy D. Hibbit ple ObservPlate 884 aFrom 24k toy Plate 08 alate Ca by Louis W adult’ male, Plate 908. C SA. Photo Airport Calsbod, San Di Photo by Troy D. Hibbs. , Plate 912. Crt a © Ko © dul male, ald anyon Davis County, Uiah, USA. Photo by Louis W, Skull Vale, Tooele County, Uta, USA. Photo by Lo Tko Cavity, Nevada, USA Ponas W. Pores tus, mole, approximately Plate 916. Coils ol o, Specimen courlesy of EI Mundo de Paste México, Mexico. Photo by Exc N. Smith, cot lesy of Manuel Varela-uls at ak Hien Plate 917. Crotalus lytic, adult mle, UTA collsc Plate 918, Citas plytictus, female, 29cm TL, UTA Plate 919, Croab Hin, Southeast Jalisco, Mente, K rom 2Akm northwest of Tapalpa, Jalisco, by Ed Cassano, Mexicoas pric pr Daring, Mico, Photo by Ed val ae Bets rice pri. Liao Grande, Durange, Proto by Jahn H. Tashan atthe Fort Wort evo Lin, Mexico. P pric subsp. adult male, UNAM- Plate 928, Crotalus price’ subsp. Same as Plate 927. NIL Vicinity of La Congoja, Mari Gracia, Aguascalientes, Mexico, Photo by Davi. Batker, courts wile, adolt, UTA 7164, Near a us, female is, Mion, Mexico, Photo by Louis W of Hui te the lange hea platIRS1455 Santa Ines, Chimalaps, Oxxaca, by Bri. Sith, Plate 959, male, approximately Plate 960, fisem TL. fQcm TL. Co Costa Rica. m adult. Mexico, Plate 964. Cvtatu 08°), elevae Phot by David ( 0 Pate 965. Cro toakom, adit; Yuratin, Plate 966°CTL, Nese USA, elevation Beis by Join Donovan and Marty Cooksey, adult. Vicinity. of a ris ins, adult, San Juan County, b USA. Patermess Utah, Us Louis W. Porras Busser en encountered througout the range Pho by Lous W.P Pate ri, adult, Huachuca Maratains, Cu ity, Arizona, USA. Photo by iments ad. Banderas ie {ion approximately 2300. Photo by Dav G. Darker atthe Dallas ZooPate 1028.inatus, Seale,Plate 1054, Chi us, female, approximate 2otem TL. Vicinity of Flor de Panga, upper USNMS 066645, Vereda de Portachuelo, vicinity of Manzanares, Meta, Colombia, elevation 10m. Tahuayo, Loreto, Peru XQ male, approximately 50cm Plate 1056. Ciccolo, female, 68cm Th, ZURC 98 TL Upper Rio Oroza, Loreto, Pet. Farenda Santa Ins, Poconé, Mato Gress, Bazil. Photo z by Ivan Sarima sally, The specimen on the right i uve ult a Mate 1059. Cn pam joven. Sea ds Mes late 100 Cli rit, approve 10cm LU Viinty oF Ente Rios, Tara, Bolivia, Photo B Pte 1088. lela womans, juvenile, ZUBC 197 Estadual, Campos do Jondio, io Paulo, Brazil, Photo Gai, Beal Phota by Nek Jonge da Silva, by rac Gece. Diet B Hares sage Plate 1061. Clels ine, male, approximately adult Pate 1063. Cony is cleats, mak TL Aguscle, Crtago, Costa Re. Photo by A 34cm TL, UTA 3 Huchuetenangt Aro Solorzano a the Serpenara Naconsl, Sa Jo Guatemala, PhoTL, UTA R i TLUTAR: Phat Plate 107 crirons, female, 34cm Tl, ZUEC sna Carina, Bri. Photo by Ivan female, approximately Plate 1080. Dj Iba Bela, $50 Paulo, Bazil, Plate 1081. Be TL Sen. Loreto, Per Photo by Otivic la. Pho TL, UTAR: Plate 1088 male, approximately Plate 1084 Aduit. Jmper Counly, Plate mor juvenile female, UTA Rr ‘uty, Tease, USA. P Barker ol ae, Pale 305. Ey sims, male TL, Plate 1096, female, 967em TL of the | ZUEC Paulo, Brazil Photo by Iwan ZUBC 1377. § s Rig de Janeito, Dex zim, Photo by fan 8 i nOdem Th, CVUCG B16. Una H That ky Paul Fed. crumbs, ul reedmL, Plate 113, Enh smal USM 6608. Vicinity of Tujshik enePlate 1124. Gancxphis Aberin Minas Ceris ra Photo by ran 87m, Plate 1128, Hotesdon Mount elmon, Cherokee County, Texas, US Photo by Plate 131, Het Florida, USA, Photo by fim Plate 1132. Flydadymasts bicincts. Senra_ da. Ne bridges Nelson Jorge da i female, approximately 70cm esa Plate 133. Hystoynases gigs, juveniles. Asuncién, Mate 134. Hyd 1s TL Mishana, 8Plate 100 dem TL. GPlate 188, Li Goi, rol sm TL, Plate 1188 rs, female, cm TL, Plate 1199 UTA R808 © Pertanbat ult ema, 94, Last e © us mae, 46cm TL We Mang rf v Ia Sera, SPlate 1199 Houston Ranch, 2.8m south on Texas Highway 1, Lln int, Tena 1sem TL, UTA — Plate 1205. Ory! FHCOAtive m, Guayas, Ecuad ok 837em TL wens, feral Country Chib, near east wu, Orange Coutly, Texas, US G. Barker UTA Plate 120 Hidalgo, Mex on County, Texas, UHorironie, Minas Gerais, Brazil. Photo by Donalmale, 48cm TL, UTA id. Finca FE Faro, -0km north of El Fal km north of El Palman, female, 420m TL, KU . nero (southeastern side of Cerro 2 5 Fortuna, You, stemala, elevation 10005, atin 180m. Photo by JR. MeCranie le Sem TE. (al ip ite rings) and male, 575em Faro, 4.0kmSanta Mari,i Sppens. Mexico Mexico, Pho Plate 1915, Ty es hypoconia, Ibiina, Paulo, Bes. PhoPate 1338 Plate 1519. Tnaste sp, female, 4A7em TL, KU 220506, Puerto Miguel, Ro Yaraps, Litto, Pera Plate 1322. 1h : i feral mm Plate 1328. Ton’ Arrstedons, So Pl 75cm 11, Ut t i raz, Photo by Otivin A. V. Marques Salle County TespproximatePlate 1340. Nena Brasil. Pho Plate 1383, Photo b Plate 1341 Plate 1345, Xewnon TL, UTA R288, Fin Plate 1348 ur 7. Racho laabal, Guatemala Plate 1350 dem TL. Finca Las Orqusensy almen Jar, i b Poul Freed Taisha, Morona-Santiga, Ecuador sitiam E Dulin, Same as Phte 1361. Pate 1363, Bots aspen, TL, El Caucho, Tubes, Peru. 7 ot locality known for B. ayer. PtPlate 1364, Bothrops sp, 110.0em TL. Bagua, Amazonas, Plate 1365. Bothnps otros, 80.0co¥TL, Pucallpa, Ueayal Por Photo by Pablo Vena Pens. Photo by Pablo Venegas.Pitvipers, Family Viperidae RATTLESNAKES Long-lived, slow-reproducing species such as rattlesnakes may, like the endangered California conde, require complete protection from man in orcer to ensure adequate annual survival for population maintenance Parker and Brown, 1974 Ralilesnakes have perhaps attracted more afenton than any ter group of snakes, They were subjects of great interest to shecorly European colonizers ofthe New World, and a number ofillusteations were published during the fist half ofthe v= fntenth century, the first perhaps being Hemanez’s Rerun afc: Noose Hisponige histor in 1623 (Klauber, 1948), Benupee and Duval (1998) summarized the importance of rat Hesakes as model organisms for diverse stadia, inccing physiological ecology life history, and behavioral ecology. The 52 species of atlesnakes currently recognized are placed into two genera, Sisrurus, which comprises 2 species, ischaracer- ized by tho presence of ine lange, symmetrical plates on the ead rots comprises about 3) species, most of which have rumerois smal scales on top of the head, especially in the frontal and. parietal regions, Crolalns nous. was previously lice inthe genus Sistarus on the basis ofits enlarged head hes (a prmiive character, but ther anatomical features ancl rncent moleclor and momphological studies suggest that i is mote properly allocated vith Crofais, with which it shares derived skeletal and hemipenial characteristics, As Klauber (1972265) pointed out, the distinction between Sistrurts and Cots based om the condition of esd scalation (ine head red ciel eal) eta pas tee definitions imply. Many species of Cras have lange head Plates on the crown, and conversely itis not unasual to find fpecimens of Sktrurus with fragmented head scales, Some se Cresta uml have sbont four larg, acizngular soles in the internasal- prefrontal region (ex, . sls, C furs, C. moles, C, psi, C sms, C. felons, se Figs. 204,212) C. mos retains the colubridlike pattern of lange head plas iypical of Sisrurs, but the parietal scales often ae par- lly frgmented. I also worth noting that in his hey to the Gpnera of pitvipers based on skull characters Klauber (1972196) thas unable 10 distinguish C. ros from olher Crofts. The lateral process ofthe squamosal of both Sistruriscattus and S iarius forms a fork having an angle of 8D degries or less Mth the longitodinal ais of the rain bono, In all specie of oii, ncucing rus, the lateral process ofthe squamosal ay be blunt or pointed, but ft forms a fork with the longi tudinal axis of the main bone, the angle formed between them is 90 degrees or more. Further, the squamosal in C.raous is rel atively short compared with species of Sistrurus and more closely resembles the bone found in certain other montane species of Crotalus such as C. lepidus and C. triseriatus (Brattstrom, 1964), Although the rattlesnakes are placed in two genera, there is, rho doubt that all are closely related and form a monophyletic ‘group. Evidence for this argument may be seen in the evolution fof a single remarkable and complicated structure not present in any other species of snake—the rattle (see Zimmerman and Pope, 1948, for a discussion of the rattle’s development and growth). All species of rattlesnakes have rattles, although several populations are unable to acquire rattle segments luring the shedding cycle ‘The rattle is used only as a defense mechanism, at least in adults (Greene, 1997). One of the theories regarding the rattle is that it evolved as a protection against being trod upon by large ungulates such as bison (Barbour, 1922, 1926; Hay, 1887a; ‘Klauber, 1940). Most of the available evidence, however, sug gests that the earliest radiation of rattlesnakes may have ‘occurred in the mountainous regions of Mexico (Gloyd, 1940; Smith, 1946b) and that the common ancestor to rattlesnakes ‘was a montane species (Klauber, 1956; Murphy, 1983a, 19830; Murphy and Ottley, 1984) and therefore probably not subject to being trampled by hooved animals, Most of the species of rat- tlesnakes that occur today in grassland habitats are not rela- tively primitive (Greene, 1997). Schuett et a. (1984) suggested that the rattle evolved as a device to enhance caudal luring, although Greene (1997), Tlebout (1997), and Moon (2001) have offered convincing arguments that refute this. The rattle may have arisen in an area that lacked abundant dry grass or leaves against which a vibrating tal alone would ‘cause a ratling sound, but instead featured rocky outcroppings fr talus slopes such as those that serve as a refuge for many species of montane rattlesnakes today (Greene, 1997), Such ‘species as C, lepidus and C., price often Sense vibrations caused by humans moving across these rocky habitats and are quick to rattle even before they are discovered and even if they are hidden beneath rocks. This behavior may warn predators such as coatimundis (Nest navi), ringtalls (Bassriseusastuts), ancl black bears (Lrsus americanus) that are known to seek small ver- tebrates for food (Greene, 1997). ‘Caudal luring is a slow, intermittent activity that would not require any special physiological modifications, whereas the til a7shaker muscles that operate aniprdator ta vibration ate spe Gilized fo sustain fest contractions for prolonged periods (Moon, 2001), Comparisons ofthe tal shaker and wunk muscles Of several species of colubrid snakes, Agkitnvon conoiris, and several ratlsnakes revealed that none of the colubrid had MUSES with significantly higher respiratory levels, even though one species (Colier constrictor vibrates it ti. The shaker muscies in both Agtishvoncnlrtric (a species hat also exhibits anpredtor tl bration) and ratlsnaee had a sg- nica higher resplratory capaciy than the trunk mines {Moon 20}, Many other raginaive notions about te fn Alon af here have been put forth, but these have now been Iargely discredited. Babcock (192), for instance, suggested that the rate served both to decay insctivoreus bids By imiatng the sound of rickets an to cll the sexes together The common belief that ralestakes cannot hea their own. rte (eg Manning, 1923) may have some basis in fac. Most Snakes, ncudingratlesnakes, hear best nthe range of 150- Got and are unable to hear sounds much over LODE but typically produce sounds over 3000Hz (Young, 1997). Some lange indivkluals of Cots, however, can produce rating sounds that have fequencis they can hear (Young and Brown, 1995, 195), Snakes are sensitive to both aiborne and vibratory stimal Young and Brown (1993, 1985 found no species-specific acoustic specializations in any of the ratlesnake species they Surveyed and conc that most ratesnakes produce brond- and noise spanning a range ftom about 2.50) to 19.0 Hi Py et al (1971) reported a range of 2000-40004, but 0 Sukeequen study has founcharange that high, The ist comprehencivetreatent ofthe struc, develop- men and origin of heat was undertaken by Garman 188, 1899), who described the rate after successive mols and com pared it with the tal spine of varius species of pitvipers: Rak flesnakes are bor witha prebutton that i shed during the frst cis exposing a button thats the fis segment ofthe rate tobe retained (oe Fig, 7), Anew rattle segment gained each time the take sheds: The number of fines that rattlesnake sheds each year is variable and depends on such things as speci, age climate, and arpount of food available. Young, fepidly growing snakes may shed four or more times a yea trheres adult sakes usualy shed no more than twice a yexr Ive nine-year study of Cc mga moch-cental Kaho in Number of rattle segments 30 10 Snout-vent length (cm) 478 Pitvipers, Viperidae which growth was carefully monitored, Diller and Wallace (2002) found that snakes shed two or three times during the first year and usually twice during the second year. Rattles become worn with age, and distal segments frequently break off, so rat Hlesnakes with long strings of segments are rare, Klaubber (1940) noted that adult rattlesnakes most frequently have 5-9 rattle segments and (Klauber 1971) stated that rattle strings of 710 segments are the most effective noisemakers Rattles of more than 12 segments are very rarely encountered under natural conditions, although Klauber (1972) reported 16 ‘segments in a specimen of C. cerasts and 23 in a C. atrox. Rat tlesnakes in captivity sometimes acquire unusually long rattles: 29 segiments have been recorded for C,liorridus (Klauibes, 1972) find 38 segments for C, viridis (Chisear and Smith, 1994). Berish (1998) showed that the number of rattle segments is positively correlated with SVL in C.adamanteus (Fig. 178). Sound is produced when two or more segments of the rattle strike each other as the tail is vibrated, Klauber (1956), Rowe and Owings (1978), and Pylka eta. (1971) described the sound of the rattle. A number of people have noted that it resembles a shorp hiss (eg, Conant, 1969; Klauber, 1971, 1972). Kinney etal (1998), using electronic analysis, found that the rattling sound ‘was acoustically distinct from the hissing sound produced in CC. widamanteus and C. alos; the rattling sound had a broader frequency (400-4,700 Hz) and higher amplitude (604B) than hissing, They hypothesized that hissing is an epiphenom- ‘enon of body inflation that is produced only during periods ‘of tapid inhalation. Fenton and Licht (1990) compared the rattling sounds of six species of rattlesnakes and found that the sound produced by all conformed to the same general pattern: medium intensity, broadband sounds with rapid onset ‘and no structural changes in frequency pattern over time. The ‘amount of variation these investigators found between species was comparable to the amount of variation found within a species, The size of the rattle varies significantly among species (Cook tal, 1994). An aroused! large rattlesnake may be heard at a dis tance of 6m or more (Klauber, 1940e), whereas the sound pro- duced by certain small species (e.g, Sistrurus mliarius, Crotalus intermedius, C. pusilus, C. steinegeri) may be difficult to detect ‘even when one is practically standing on the snake. When the interspecific differences in SVL are’ removed by allometric 180 Fig. 178. Number of rattle segments (ndieating numn- ber of shedk) versus. SVI_ in Cros adorns. ‘Adapted from Bersh, 19983553 (6 2.scaling, however, the rattling sounds of 21 species of rat- Aisnakes become indistinguishable from one another; only S. silaran produces a comparatively quieter and higherpitched tale than the other ratlesnakes (Cook et al, 1994), The acoustics of rattling in C. organs oregnus were investigated by Young and Brown (1995), who looked at variables including baad size, rattle size, ratle segment length, and different env ronmental conditions. They found that removing the distal rate segments didnot sigeificantly: change the dominant ‘maximum and minimum frequencies of the sound produced bythe rattle and concluded that the size of the proximal rattle segment is the most important feature determining the sound produced by the rattle Sisk and Jackson (1997) suggested that the ability to retain ratle segments—and thus to evolve a ratle—was contingent on the prior development ofthe longi tudinal bilobing of the terminal scale. They suggested two hypotheses forthe inital functional advantage of such bilobing the luring effectiveness hypothesis, whereby bilobing would enhance Visual attractiveness to potential prey; and the dual contact hypothesis, in which aposematic sound quality would ‘be enbuanced during tail vibration. When both hypotheses were cperimentally tested, the authors could find no support for citer although they suggested that othor aspects of the dual canact hypothesis should be further investigated Kissner et al. (1997) investigated variables including sex, reproductive status, body size, and body temperature in rela- tion to eating behavior in Croalus 2 aris. Two groups of snakes allowed intruders to come significantly closer before Ieginning to rate: gravid femalds and small snakes. The authors hypothesized that these two grOups depend on exypsis toa greater extent than other snakes because they are less able to escape prelators and because small snakes may be more ‘ayptic than larger snakes, providing them with an advantage Most analyses did not seem to show a consistent relationship between body temperature and rattling distance, but ool sravid females did allow intruders to approach more closely tan warmer gravid females. ‘The speed with which rattlesnakes vibrate ther ols is fem- perature dependent (Chadwick and Rahn, 1954; Klauber, 1971; Marin and Boyby, 1972). Chadwick and Rahn (1954 found that the lationship of rattling frequency (measured as cycles per rina) to temperature in C. 0 viridis was linear between 5 and 40°C, and Martin and Bagby (1972) found thot the relationship tenween these variables in C.atrox was linear between 16 and 52°C, which corresponds largely to the temperature range at Thich that sake is active, At temperatures above 34°C rattling became spasmodic, and between 3 and 15°C the rattle fe- quency dropped precipitously. At 155°C ratlesnakes typically Have vibration speeds of about 35 eyeles per second, but at SUC the vibration speed increases to about 85 eyeles per second (Klauber 1971) ‘ix muscles make up what is often referred to asthe shaker complex In 9 recently skinned rattlesnake these muscles are sully evident because they are a darker red than adjacent sheleal muscle. Studies on C. horridus, C. ruber (Clark and Schultz, 1980), and C. atox (Shaetfer et al, 1996) determined tbat tail shaker muscles have large numbers of small myofibrils, lange deposits of glveogen, and an extremely high volume of sorcoplasmic reticulum, which together minimize activation, ‘contraction, and relaxation times. A high volume density of mitochondria in this muscle maximizes ATP resynthesis. Shaker muscle differs from other skeletal muscle in that myofibrils do not entirely fill the fibers and the myofibrils are highly branched, giving the fibers a mottled appearance (Schultz eta, 1980}, Conley and Lindt (1995) found that the tail shaker _muscles have a reduced volume density of myofiber and require minimal ATP for contraction. Thus, the tail shaker muscleis spe- ialized to produce a sustained high-frequency warning sound. at a cost that is among the lowest found in striated muscles in vertebrates (Conley and Lindstedt, 199). The shaker muscles lf C. horridus and C. ruber can contract at arate of about 50 per second for at least three hours (Clark and Schultz, 1980), Kemper et al. (2001) showed that glycolysis in tail shaker ‘muscles provides a high andl sustainable ATP supply along with ‘oxidative phosphorylation without muscle fatigue. The size and structure of the rattle may vary considerably ‘among, populations of the same species and between closely related species. For example, members of the island population ‘fC. catalinensis lose the developing rattle segment each time they shed, and thus retain only a shrunken rattle matrix and no rattle or only a single basal segment, Crotalus ruber lorenzoensis from Isla San Lorenzo Sur also has this tencleney, and about half ‘of the individuals of this population have lost their rattles. In contrast, C ruber individuals, the mainland relatives of C. catal- nensis and C. ruber forenzoensis, have large, well-formed rattles, The rattle of Sistrurus catenatus is lange and robust, whereas §, miliaris has one ofthe tiniest rattles found among rattlesnakes, ‘The size of the rattle varies significantly among isolated popu- lations of Crotalus raous (Fig. 179), with the comparatively small rattle of the Guerreran C, nus exiguus being diagnostic for that population (Campbell and Armstrong, 1979) Besides the rattle, rattlesnakes use hissing, body postu scent glands, and extended protraction ofthe tongue as warning signals (Klauber, 1972). A few species may use an open-mouth threat or mave the head in a provocative way. Crotalus molessus is one of the few rattlesnakes that we have observed to engage n an open-mouth threat, a behavior also noted in this species by Armstrong and Murphy (1978), We have been able to elicit this defense only im a few snakes from southern populations. Armstrong and Murphy (1978) also reported a single specimen ff C. polystictus that exhibited an open-mouth threat posture. Marchisin (1978) reported observing individuals of Sistrurus tilarius harbour’ in the field and laboratory bobbing the head vertically while they rattled. He suggested that the bobbing serves as a visual warning signal to potential predators. In most species of rattlesnakes males grow to greater lengths than females, Several interesting exceptions are the sidewinder, C. cerastes (Klauber, 1972), and the pyginy rattlesnake, Sistrcrus rilarius (Bishop et al, 1996). Porras (pers. comm, 2002) indi- cated that Crotalus oreganus concolor females also exceed males in size (but see Ashion, 2001, for a conflicting view). Klauber (1937) found that males of many species of rattlesnakes were slightly larger (1% of body length) than females at birth, and that the sexual dimorphism in body length increased with age. Klauber (1997) provided information on a sample of 858 C viridis collected from denning sites near Plattesville, Colorado. ‘This series is unusual because ofits size and because the snakes ‘were collected with no discrimination for size. Figure 180 sum- -marizes the sizes and sexes represented in this sample. This his- togram is strongly bimodal, with juveniles (25.0-34.9cm) and adults (550-79.9em) being the most common, The largest size categories are composed entirely of males as 2 result of sexual dimorphism, and the percentage of males making up the larger size categories is higher for the same reason. CCrotals atrox is sexually dimorphic in size throughout its range. Beaupre et al. (1998) showed significant sexual size Rattlesnakes 479Proximal rattle segment (mm) Fig. 179. Regression of dorsoventral width of proximal fale segment on tail length for males of the thre sb- species of Cals es. Closed circles = rus, squares reas, oper cies = exis. The slope and el wall of the tegnesion for rrous are signin die ferent (P= (10D) fom those for Daves and exgus, but the regressions of brummeus and eiguos are not lg a See ot e 7? neanydifleent Tram ead other Adapted. om Tail length (mm) males Campa and Armstrong, 1979307 (ig 3 os 2 2 i & . s be g a 5 fer LL 2 a Fig, 180, Size and sex distributions for a sample of B58 wali ae wee] [100 10, individuals (459 males and 399 females) from a snake Se sian kn Rulon of Cras crits near Patri ES SS SES SG GS GG BB BGG BE Colorado, USA, gure a hel of columns indicate Size class (mm) dlimorphism in a population of C. atrox from central Arizona (Fig. 181), There was no significant gender difference in size at birth, and juvenile growth rates were similar. The divergence in size between males ancl females occured after sexual maturity ‘was achieved. Certain rattlesnakes aggregate in huge numbers in dens during the fall (Gregory, 1984; Sexton et al,, 1992). Hall (1929) found about 50 rattlesnakes aroundl or near a denning site near Ely in eastezn Nevada, He identified these as Crotalus confuen- fs kelly (=C. scutulatis), but the geographic locality indicates that they were C. oegamus lutsus (L. Porras, pers. comm). Hall also eeported that several persons had reportedly killed 139 rat- tlesnakes at this site, several days prior to his visit, and the 480 Pitvipers, Viperidae percentage of males in each size dass. Adapted from Klouber, 198720 (ig 4). number of empty gun shells and dead snakes that he found ‘scemest fo confirm that figure, When this den was first discov- fered, during the autumn about five years before Hall’s visit, large numbers of rattlesnakes were purportedly seen coiled together into balls around the edge of the entrance. Graf et al. (1938) reported that 97 C. oreganus oreganus were Killed in April 1837 and another 27 in 1997 at a denning site on the western side of the Cascade Mountains in Oregon. These ‘authors also reported that more than 100 rattlesnakes were Killed by a crew blasting a road through the region. Bushar etal. (1998) found that individual C.horrius from the same denning site usually were more closely related to each fother than to snakes from nearby hibernacula, In general,2% 2 B6 a § . 0 & Bein, sie dances fs mie mais = alpnereeh pemtrtnert omar ‘aplure, Note that males sehieve a lager SVI. than do Females. Adapted fom Beaupre ot aly 1998612 (0g 1. genetic distance inezeased and gene flow decreased as geo- graphic distance increased between hibernacula, although Certain structural habitat features such as location of suitable basking locations tended to indluence genetic variation in ways not strictly corvelated with distance ‘Aden discovered in Toosle County Utah in 140 was the focus of studies by Woodbury and collaboriters (1951). During the Course of ther studies they marked 930 rattlesnakes and had W080 recaptures, No other dens were found-within a 3.2m radius. One marked snake was captured about 2km from the den, Many more adults than juveniles enfered the dens than tmght be expected ifall snakes were equally ikely tousetheden, eos (n Woodbury etal, 1951) slated that an ar temperature of 6° 15.6%) adjacent tothe enteanceof the cen was the thresh- dl temperature at which denning C.oreganus (asus) became active and came tothe surface. In most ofthe 11 winter seasons ofthe study (1939-1950), the numberof males coming tothe den facexceeded the number of females Qulian, i» Woodbury et al. 1851). Only in the winter of 1940-1941 did females outnumber males (95 females versus 85 males). Over the total period of the Stuy, 1,179 males and 803 females were recorded from the den Julian (in Woodbury etal, 1951) suggested that sex ratios were about equal forthe first three years of life in this population of taltlesnakes but began to diverge at about the time of sexta maturity owing to differential survival rates ofthe sexes. Gliss- sreyer in Woodbury etal, 1951) reported that females arrived at the den in the fall with enfarged ovarian eggson alternate years, suggesting biennial reproduction; 125-66.7% of the females ‘were gravid in particular years, with an average of 47% forthe ML seasons, Ovulation occurred in May or June followed by fer ization, and young were bor in August or September. Females tarred 2-9 =5.5) eggs Hirth (19660), who also studied snakes from this den, found the smallest mature male to be 65cm in SVLz the two salest sexually mature females were 54 and 56cm in SVL Heyrend and Call (in Woodbury et al. 1951) found that most ratlesnakesshed tice the frst ull active season after their birth, ater which time there was an increasing tendency for snakes to molt only onceeach season. Itwas unustaltofind more than six to nine rattle segments on a snake. ‘Afollow-up study at this den was undertaken from 1969 to 1973, at which time the population had declined to 12-17 rat- mm Holos al [= Females > o 0 © 7 9 9 io 10 Snout-vent length (em) Hlesnakes (Parker and Brawn, 1974). Overwintering, mortality amounted to 3.6% for the four winters, whereas summer mor- tality averaged 17.8% per year. The overwintering weight losses amounted to 4-9%, with females losing more weight than ‘males. No snakes younger than five years old were found, and the age structure had shifted toward older age groups. Based ‘on a simulated life fable and demographic traits of rattlesnakes, Parker and Brown (1074) estimated that this long-studied pop- tlation was near extinction at the time of their study, and a brush fire in the area on 17 June 1974 may have extirpated the fow remaining individuals Weight loss during hibernation is no doubt responsible for considerable mortality, especially of juveniles following unta- vorable years. Klauber (1972) estimated that adult C. ovis con average lost about 4% of their weight during hibernation while juveniles lost about 20%, Hirth (1966b) found that weight Toss in C. 0 lvtsus from a den in Utah was about 6.3% and 8.5% in adult mates and females, respectively, and 25.5% in juveniles (less than 31cm), The mean number of days spent overwinter- ing in the den was 198 for juveniles and 219 for adults. Hirth estimated the overwintering mortality for rattlesnakes at this site to be about 34% (based on the recapture of 23 of 35 snakes), but some surviving snakes may not have been recaptured, 0 this figure may be too high. ‘An estimated 300 C. oreganus (Iutosus) were killed in one den. in Tintie Canyon, Juab County, Utah, in about 1937 (Woodbury and Hansen, 1950), but the den was still active in September 1949 when it was encircled by a screen wire fence with several traps to catch snakes coming to it. Five species of snakes used this den, including Crotalus oeganus Intosus, Masticophis tnenia fs, Pitophis cotenifer, Lampropeltstrizngulumt, and Tharmophis “oninoites (=, elegans vagrans, fide L. Porras). Of these, C. ore= ‘gums accounted for about 54% of the total catch (69 of 127, snakes) ‘At a den site discovered in 1952 in Tooele County, Utah, ‘Wooxlbury and Parker (1958) attempted to trap the entire den population. During the spring and fall of 1953 four C.o, Iutosus, two Pituophis calenfer, and 85 Masticophis taeniatus were collected. Parker and Brown (1973) studied a series of snake hibernac: ula in northern Utah and found that Coluber constrictor, Masti: Rattlesnakes 481cophis tacit, Pitwophiseatnifer, and Crotalus 0 llosus over Wintered in these dens, but that ratlesnakes were restricted to particular dens. One of these dens had been studied by earlier workers and provided information on long-term changes. A total of 252 Coregonus llosus were observed at this site in 1950, 55 were sean in 1966, and only 12 were observed in 1972. Killing by humans is almost certainly the primary reason for the pre cipitous dectne. thas been known for some time that snakes require heat to digest prey. With the advent ofthe fist cold snap ofthe fall, most snakes appear either to ease feeding orto feed only infre= quently thereafter After returning to dens in the fll, snakes continue for several weeks to emerge on warm days ta bask prior o entering the den for a quiescent period of several to many months I has been suggested that fasting i necessary prior ta hibernation to void the gut of food that might putzely Causing harm or even death to the snake (Gregory, 1982} ‘Macartney’s (1989) observation of dried snake seats foun in great abundance around the entrances of rattlesnake dens sup- ports the notion that these snakes void most of the digestive tract before cold weather permanently sets in. Diller and Wallace (1984) found that adult female C. o. ov gas fom Idaho emerges irom hibernation with enlarging ova, ‘vlad rom mid-May’ to-mid-June, and gave bith aver about 4 Sl-day period boginning in early September. The mean litter size was 55, and the number of young, was strongly cortdated ‘with female body size. In the several populations studied, the percentage of females reproducing annually varied from 54 to 27% wh he fequeney of repro pinay infuenced by nutrition. These authors found that males Jonger than 52cm in SVL and having four or more rattle segménts were mature, tout most mature females were moe than 35cm in SVL and had five oF more rattle segments In Sweetwater County, Wyoming, pregnant females moved only a short distance toa rookery after emerging from hiberna tion,and they remained there throughout gestation without for- aging or mating (Ashton and Patton, 2001. In this population tlspring were born between 20 August and 18 November, with 4 mean liter size of 17, a mean offspring SVL of 19.3em, and a mean offspring mass of Sg; females appeared te reprecuce every two or tree years (Ashton and Patton, 2001) ‘A population of C.v. viridis near the northern limits of the species’ range was active from April to October, and about half of the adult females were gravid each summer Secoy, 1984). Gravid females of this population were signifi- cantly more ative during ll seasons Gannon and Secoy, 1985) In south-central Wyoming, Duvall et al. (1985) found that female C. 0. virds gathered in communal bitthing, rookeries where they gave bith by early September. Males ofthis popu lation made lengthy vernal foraging, migrations in excess of 5km from the denning site; both males and. females were attracted to sites containing live deer mice (Duval etal 1985, 1990}. King and Duvall (1990) found that both males and females foraged for food during about the frst haf ofthe 35+ month active period in Wyoming: during the second half of the active season the attention of females continued tobe on seareh- for prey, but males searched for both food anc mates (King, and Duvall, 190). ‘The ageat which rattle snakes reach maturity docs notappeat to be strongly correlated with size or geosraphic location, although no doubt both factors impose some constraints. Cro (ols herridus, one of the largest rattlesnakes, requires four to seven. years across its range to. mature (Brown, 1991). A non and 482 Pitvipers, Viperidae moderate size satlesnake, C.oregarus orgs, matures in five toseven years in the northern portion of is range (Macartacy etal, 1890) and takes only three yearsin more southerly regions (Diller and Wallace, 1984; Fitch, 1949). Most females ina pop- lation of C: eres ror north-central Idaho reached sexual ‘maturity during their fourth stummer, but males in this popu- lation matured more quickly sometimes as early as thei second summer (Diller and Wallace, 2002). Most adult females from this Idaho population reproduce biennially, but in some instances may reproduce during two consecutive years, presumably when prey densities provide sufficient energy for annual repo> duction (Diller and Wallace, 2002). Crotalus avox, another large snake, achieves maturity in thee to four yeats (Beaupre etal, 19965; Fitch and Pisani, 1993). Secor (19940) estimated that C «erases in the eastern Mohave Desert are mature af thee years of age Finally, the relatively small rock ratlesnake,C.lepidus lepies, found in relatively mill climates, requires seven years to reach maturity (Beaupre, 1993). Females may travel cnsiderable distances fom denning sites before giving birth (Duvall etal, 1985; Galligan and Dunson, 197; Reinert and Koxrich, 1982) How young snakes make thet way tothe denning site forthe fist ime is not well understood; it fs possible that first-year snakes utilize different overwinter ing sites (Hieth, 19569; Swanson, 1952; Woodbury ancl Hansen, 1950), In the case of Choris and C. viridis, which have been relatively well studied, at least some neonates use the same winter rehugia as adults use (Duvall et al, 1985; Reinert and Zappalort, 1988) raising the question of how these hibernac- tla are located by neonates. Both laboratory angi fel observ2- tions on C.laridus strongly suggest that young, C, horidus use chemical cues to tall adults to denning sites (Brown and Maclean, 1988; Reinert and Zappalorti, 188b)-and this method seems probable for other denning species as well The physiological ecology of ratlesnakes isa very new field, ‘uta few recent studies offer intriguing glimpses into the world of rattlesnakes that have a significant bearing on our under: standing oftheir ecology and evolution. For example, Beaupre (1996) studied metabolic rate and water flux in two populations ‘of mottled rock rattlesnakes (Crotalus fepidus) in. Big, Bend National Park, Texas, and found that one poptiltion had an annual energy budget about half the magnitude of that ofthe ‘ther population, indicating a considerable difference in food resources. Not surprisingly, snakes from the low-energy: budget, food-limited popiation hal lower growth rates and attained smaller adult body sizes thon snakes from the cther population. Snakes from the lower, hotter laity main tained significantly higher body temperatures as a result of environmental thermal constraints (Beaupre, 195a) In August, the mean body temperatuce of the hot-loeality snakes was about 1 to almost 5°C higher during diferent times of the day (Fig. 182) Body size varies dramatically. from species to. species ‘Members of the genus Sistrunsand ofthe species Crotalis rans ‘ers, C.intermetins, C. rsritus, and C. wile usally reach only about 7Dem in TL and weigh less than 05k, while members ofthe species C. alas, C. fusca, C. durin, C.har- vids, C. molossus, and C. simas often exceed 10cm in TL and may weigh in excess of 10k, The adult C.adntens isa truly formidable animal, approaching oF exceeding 250 cm in TLand reaching in excess of 3Dlbs (136k), the greatest weight of any venomous snake (Klauber, 1971). Klauber (1938), who measured head length in rattlesnakes in relation to SVL, noted that only C. corastes exhibits sexualMean Body Temperature (°C) ig. 182. Daily body temperatures (mean +255) of to populations of Crofas pid in. Big Bend National Fath, Teaay USA. The Boquilas Canyon (BOQ) popu Iain lives ta elatively low elevation; the Grapevine ° Hills (GVH) population occurs ata higher elevation ‘Adap fomn Besse, 1995160 ig. Be er ioe e es 5 10 6 » Hour of Day While Crotalus cermsts is well adapted to occupy sandy areas of the Mohave Desert, however, some species of rattlesnakes: thrive in Mexican cloud forests; two more different habitats can scarcely be imagined. Many species live in relatively flat low lands or rolling prairies. Some species are found in marshes (Gistrurus catonatus) oF forests. McCoy (1984) placed S,cafenatus in his “semiaquatic” ecological category, although it should be noted that this category included species living in marshes, sedge mals, and other habitats peripheral to watercourses and Jagunas; this species is certainly not as closely associated with ‘water as the natricines, which are truly semiaquatic. Klauber (1872) voiced the opinion that C polystictus might be somewhat ‘aquatic based on the field notes of Paul Ruthling, who collected this species near Lake Chapala in Jalisco, Mexico, Armstrong and Murphy (1979) suggested that while C. polystcius may inhabit lake margins that are subject to flooding after heavy rains, this species is not semiaquatic and is solely an inhabitant ff montane meadows, and flatlands. We have most often seen . polysticius in the high, open grassy valleys and the rocky ‘mesquite-grasslands of the southern Mexican Plateau. We hhave encountered individuals a few meters from slow-moving, meandering steams, but never in what might be considered an Aguatic situation. We might be inclined to consider Rutling’s observations aberrant were it not for an even earlier, often foverlooked, independent observation, also made near Lake ‘Chapala, that clearly describes the aquatic habits of this species (Beebe, 1905; see the C. palystctus account) ‘The overall ground color of rattlesnakes is often closely asso- ciated with their habitats. For example, Crotalus intermedius individuals from the deserts of Puebla are pale gray with dis tinct darker brownish gray blotches, snakes from the open pine- tak forests of Oaxaca and Guerrero have @ somewhat darker ‘ground color although the dorsal blotches are still conspicuous, fand snakes from the cloud Forest of these states are usually dark Rattlesnakes 483_gray to almost black and dorsal blotches become obfuscated in adults. Crotalus ravus specimens from dry habitats in Puebla and Oaxaca likewise have a distinctly pale ground color with well: defined blotches; those in the highlands of Guerrero usually have a somewhat darker ground color; and those in the cloud forest near Totontepec, Oaxaca, and the high-elevation fir harclwood forest near Huitzilac are mostly dark brown. Several populations of ratlesnakes that live in dark lava fields in the southwestern United States are melanistic, including C, atrox from the Peco Armendariz lava field of New Mexico (Best and James, 1984); C. mitchell from the Pisgah lava flow in southern California (Norris, 1967); and C. molossus in New Mexico from the Tularosa malpais (Lewis, 1939), the Afton lava flows (Prieto and Jacobson, 1968), and the Pesiro Armenclariz lava field (Best and James, 1984), Rattlesnakes are good swimmers, and some will freely enter ‘water, Dunn (1913b) captured a specimen of C. horridus swim= ‘ming, in the Shenandoah River of Virginia, and Frey (1996) observed a C. lpitus sitting on a ledge about 50cm under the Water in a small pool of the South Fork of Negrito Creek in Catron County, New Mexico. Frey suggested that the snake may have been foraging for fish, or altematively that its aquatic behavior was associated with thermoregulation (the air tem= perature was about 35°C, while the water temperature was about 16°C), Some species of rattlesnakes have no trouble breaching narrow saltwater barriers. Many Atlantic and Gulfcoast islands. off the southern United States are inhabited by rattlesnakes, especially C. adamanteus-and C. atrox; Catavrba and South Bass Islands (Ottawa County) itv-Lake Erie are known to be popu- lated by C. horridus (Walker, 1931); and islands populated by rattlesnakes are especially characteristic of the Gulf of Califor- nia (Campbell and Lamar, 1989; Grismer, 1999, 1999, 2002; Murphy and Ottley, 1984; see Map 92), where some populations hhave become sufficiently differentiated from their mainland ancestors to be regarded as distinct species. Few islands off the Pacific coast of North America are inhabited by rattlesnakes, undoubtedly because of the cold ocean currents that separate these islands from the mainland: however, rattlesnakes have managed to reach two Pacific islands: Cedros Island (C. ruber) and South Coronado Island (C. organs), In terms of their elevational distribution, rattlesnakes are among the most successful of the New World snakes, having adapted to climatological conditions ranging from sea level to alpine environments, In California, C. oneganus feller is known. from 3,293m on San Jacinto Peak (Ewan, 1932) and reaches about 3,350m elsewhere in the state (Klauber, 1934). A lange radiation of small montane species occurred in Mexico and the southwestern United States. One of these species, C. triseriatus, is known from 457m (Klauber, 1972), almost at the snow line, fon some of the great volcanos of the Transverse Voleanic Cordillera. Although reptile life would seem most unlikely at such high, cold elevations, on sunny days the dark volcanic rocks reach surprisingly high temperatures (Swan, 1952; see Fig. 183), It fs no coincidence that montane rattlesnakes are associ ated with rocky outcroppings, which both provide radiant heat and afford shelter to the snakes’ principal prey, izards of the igenus Sceloporus, Species that exhibit particularly wide eleva- tional distributions include C. basilscus, C. horids, C,oregans, C. simus, and C.tofonaeus. A few species (C. adamant, C. trans versus, and the species inhabiting the islands in the Gulf of California) occupy relatively narrow elevational zones Rattlesnakes do hybridize in nature, but rarely. Based on mor= phological evidence, Campbell et al, (1989a) discussed an apparent hybrid between Crotalus lpidus and C. wlan from. the Peloncillo Mountains of New Mexico, This hybrid was similar to C. feius in the mumber of scales in the internasal- prefrontal region, the number of loreals per side, the nature of the subocular-supralabial contact, the supralabial pattern, the postocular stripe, and juvenile tail coloration; it resembled C. ‘lla in having an undivided upper preocular; in the number ‘of ventrals, subcaudals, and dorsal scale rows at midbody: and in the body pattern. It was intermediate in many characteristics, Perkins (1951) reported several instances of hybrid ratile- snakes produced in captive conditions. In one case a male Crotalus oreganus helleri mated with a female C. ruber ruber, pro- ducing 9 viable offspring (Klauber, 1956), Another case involved a female C. durissus unicolor that mated with a male C. scutila- tus scutulatus and produced 4 young, These young eventually reached maturity and bred among themselves, producing, 2 litter that contained 4 viable young. Hybrids between C. satamantews and C, horridus have been foundl in South Carolina and Florida (Pl. 758). We have seen photographs of naturally ‘occurring hybrids between C.scutulatus and C. molossus in Texas and Arizona, between C. scutulatus and C. atrox in Texas, and between C. viridis and alrox in Texas. Based! on various venom characteristics, including the pres- ence of Mojave toxins, Glenn and Straight (1990) suggested that hybridization occurs between C, viridis ori and C. scutulatus scutulatus in southwestern New Mexico. Murphy and Crabtree pes 2. f i a a te F 7 E —a . it ka Gane [Sn Fane Goorniaht aT 2S 45 67 8 811MM 1 1017 We 19.20 22 2D Hour 481 Pitvipers, Viperidae Fig. 183. typical day a 120m on Volcin Orizabs, “Mesico. Adapted from Sion, 1952108 (ig.(2988) used allozyme data to identify a hybrid between Crotalus scutulatus and C. viridis viridis that had been collected in “Hudspeth County, Texas. A female Crotalus seutulatus from Kern County, California, and a male C. oregamus oregumus from Contra Costa County; California, that were maintained together were ‘observed eopulating on 10 May 1952 In late September or early (October the female gave birth to 12 young. Six of these young, had a color pattern similar to C. scutulotus with a greenish ‘gmund color and blotch borders of unicolored pale scales, 1 Individual from this litter closely resembed C.oreganus oreganus Key to the Species of Rattlesnakes" Jn color pattern, and the remaining young were somewhat intermediate between the parental species for color pattern, favoring one or the other, Bailey (1942) reported what he thought was an intergeneric hybrid between Sisfrurus coleatus ‘atenatus and Crotalus horridus from Towa, Other terrestrial snakes that are sometimes confused. with rattlesnakes include members of the genera Elapie (Pls, 1090-1084), Heterodon (Pls. 1127-1131), Hypsiglena (Pl. 1137), Lampropetis (Pl. 1142), Pituephis Pls. 1243 and 1284), and. ‘Trimorphadon (Pls, 1327-1329), 1. Top of head with fewer than 12 lange, symmetrically arranged plates including paired internasals, paired prefrontals, and a single frontal 2 Top of head with more than 12 smaller, asymmetrically arranged seales, including several in the frontal area and numerous scales in the parietal region 4 ‘Canthus angular with lateral edge of canthal scale bearing moderate ridge, not curved downward on side of head; rostral high, truncate at top; body blotches usually as wide as or wider than long, 3 Canthus rounded and lateral edge of canthal scale cu to a point as it curves posteriarly between internasal 959), ing downward on side of head; rostral low, tapering Body blocs usually longer tan wide (Ps. 93 Crotalus 3, Upper preculor usually in contact with postnasal, anterior suboclar ually contacts supralabials 4 and 5 (Ps. 1011-1017) ‘Sisrurs teats Upper preocular not in contact with postnasal, anterior subocular usually contacts supralabials 3 and 4 (P's. 1018-1023) 4. Lateral edges of supracculars raise into at, pointed, hornlike process (Pls. 791-794) Sistrurus liars Croats cerastes Lateral edges of eopeSocals nt extended into pointed, hore process 3 5, Fewer than 40 subcoudal les in males and fever than 35 in female 6 Mare than 40 subudal sales in ales ane more than 35 n foals u 6. Tip of snot and anterior portion of cath asd into sharp igs rostral and mana fen wih a median vertical pale line (Pls. 98-1010) (Crofalis ean Tip of snout and anterior portion of canthus not aise int sharp ridge rostral and mental not marked eth a ‘mean vertical pale line ? 17, Prenasals separate from rostral by small, often granular, scales; upper preoculars often divided horizontally ‘oe vertically, both (Pls. 864-875) (Crt ithe Prenasals contacting the rostral upper preculars usually not divide oy, ifso, divided only vertically with ante shor portion of scale conspicuously higher than posterior section and curved over the canthus in front of the supraoeular : ieee 5, Aseries of 35 or more crosshands formed of dark dots ona paler background, dorsoventral width of proximal rattle segment les than 25 times head length (Pls. 958 and 959) Crotalus tigris Dorsal body blotches shaped as diamonds, hexagons, rectangles, ovals, oF ellipses, of if crossbands present, these not made up of conspicuous dots; dorsoventral width of proximal rattle segment more than 2.5 times, heal Tenth $ 9, Anterior subocular contacts 1 or more (usually 2) supralabials 0 Anterior subocular not contacting any supralabi v7 10, Supraaculars transversely crossed by a thin, black-bordered pale line; a distinctly outlined round or eval dark blotch below the eye; dorsal body pattern of longitudinal ellipses (Pls. 915-918) ‘Crotalus plystictus Supraoculars not transversely crossed by a thin, black bordered pale line; no distinctly outlined round or oval dark blotch below the eye; dorsal body pattern nat of longitudinal ellipses 0 11, Midbody scale rows plus supraabials on both sides total 41 or fewer «.. . 2 Midboxly scale rows plus supralabals on both sides total 42 or more “ 12. Loreal in contact with 1 or more supralabals (Ps. 815-€09) Crotalus intermedius Loreal not in contact with any supralabial B 13, Lower preceular passes into facil ptand fal to make contact or oly very narowy 20) with the loreal body pattern of undivided dorsal crssbands (Pls. 978-980) (Coals trascersis Lower pruocular passes above the facial pit and makes contact with the loreal; body pattern usually ‘of paired spots (ome of whieh may be irtegularly connected across the dorsal midline) (Pl 19-528), . ‘Crotalus price, Rattlesnakes 485486 1. 15 16. ”. 2 a 24, (Only 2 scales in prefrontal area (Pls 929 and $30) Croats pusits More than 2 scales in prefrontal area 15 Upper proocular usually dvicied vertically, the anterior section being higher than the posterior and curved over the canthus in font of the supraocular; dasal body batches (fewer than 25) occupy ess longitudinal space along midline than interspaces (Pls. 841-863) .. (Crotalus pide Upper preocular not divided vertically of, if divided, the anterior section not higher than the posterior {and not curved over the canthus in front of the supraocular; dorsal body botches (25 or more) occupy more longitudinal space along midline than interspaces. 16 Rattle-finge scales usually 10; rattle relatively large, in snakes exceeding 40cm in TL the dorsoventral width ‘of the proximal ratte segment is more than 1 ofthe TL (Pls. 756-772) ‘Crotalus Rattle-finge scales usually 8; rattle relatively small, in snakes exceeding 40m in TL the dorsoventral width of, the proximat rattle segment is less than 1% ofthe TL (Pls. 981-987) Croats triseriatus ‘More than 2 scales between the nasals in contact with rostral (usually termed internasals) (PIs, 889-914, 988-997) ‘Cotes oregon, C vis (see species accounts) ‘Two scales (internasals) between nasals in contact with rostral 18 Upper peeocular usually divided vertically, the anterior section being higher than the posterior and curved over the canthus in front ofthe supraccular; dorsal body botches occupy less longitudinal space along midline than interspaces (Pls. 11-863) Crotals lids Upper proocular not divided vertically of, if divided, the anterior section not higher than the posterior and not carved aver the canthus in font of the supraocular; dorsal hody botches oceupy more longitudinal space along, ‘midline than interspaces 19 ‘Ventral 163 oF fewer 2 Ventas 164 oF more : 2 Haieeige scaled call 10, te iat latin oaks waving scr in Tt dersovertial wth ‘ofthe proximal rattle segment is more than 1% ofthe TL (Pls. 756-772) Cotas agus Rattle-fringe scales usualy 8 rattle relatively small in snakes exceeding 401m in TL the dorsoventral width of the proximal ratle segment i less than I of the TL. (Pls. 81-887) Cros triseritus ‘Tail rings or bands highly contrasting and conspicuous, alternating white or whitish gray with dark beown or black 2 “Tall black or datk gray; rings or bands, if present, only maderately evident because of lack of contrast between dark rings or bands and ground color 28 Kale matrix shrunken, o single rte segment present, no loose ate sogments (P78 730) ‘Crotalus eatatinenss Rade ate otal staly worl lobo ate sare present (xcept in very small juveniles which may have only a button) 23 [asta agar berets abo eal b ogress th plait innit ber Wf nln Bates ssupraoculats rarely less than 4; no flat erescentric scale bordering each supraocular posteriorly 2 Dark tail rings shorter than pale interspaces; pale postocular stripe if present, extends posteriorly above ictus; ‘minimum number of scales between supracculars usually 2; usually a flat crescentric scale bordering each supraccular on the posteromedial side (Pls, 949-955) Crotalus scutultus A vertical pale bar on prenasal and first supralabial; postocular pale stripe extending from posterior edge of «eye diagonally to point above or behind angle of mouth (Pls. 782-785) rveve CMMs adaraanteus No vert pale bar an prensa an fis suprlbial posto ripe extending fom posterior age of sliagonally downward t0 ultimate or penultimate supralabials Fist par of inalabials wsully transversely divided; dorsal blotches not conspicuously marked with back punctat 2% First pair of infralabials usually not transversely divided; dorsal blotches conspicuously punctated with dark OES eves 2 say oe sae prnasa sly contacting fstsprlbia data ings sly not neropted laterally (Pls. 940, 941, 945-948) ,. ‘Crotalus raber Intergenial scales wsually present; prenasal usually not contacting first supralabial; dark til rings often inter: rupted laterally (Cedeos Island Ps. 922-B44) «ss... ‘Cros ruber Upper preccular usually in contact with postnasa, oF such contact prevented by an upper loreal (Pls. 773~ ps rola arox Upper preaculae usually not in contact with postnasal, and no upper loreal present (Pls, 970 and 71) Crotalus tortugesis Pattem on bodly of black chevton-shaped erossbands (Pls. 829-834) Cotas horrid PPattom on body of diamond-shaped, squarish, or roundish blotches. 2» Internasal prefrontal region covered with 13-25 scales; crown and frontal arva covered with mostly rough, ridged, and knobby scales (Pl. 825-828) ereeveeneneseesess CRORIS enYD Pitvipers, ViperidaeTntemasal-prefrntal region usually covered with 12. or fewer sealeserown anc fromal area covered with mostly flat and smooth sales - ei aD 30. A pai of distinct dorsolateral stripes on the neck 1-3 scales wide, separated by pale middorsal interspace 2-3 seiles wide, and extending 1-1 head lengths behing head before reaching the first dorsal Blotch soo... 31 [No paired dark dorsolateral stipes on neck os if present, irregular and extending posterior less than 1 head length behind head 2 431, Most dorsal diamond-shaped blotches separate along dorsal midline and not extending down sides as eros bands: paravertebral stripes on neck bordered laterally by a pale scale row that isnot conspicuously paler than adjacent area below (Pl. 956-966) i cecsesnn saves CPM sims Dorsal diamond-shaped blotches mostly in contact along dorsal midline; paravertebral stripes on neck ‘bordered laterally by a pale scale rove conspicuously contrasting with next scale row belovy on sides of neck (Ps, 795-824), Crotalus duyissus 32. A pale-bordered, black of dark brown bar erasing the head between the anterior points of the supraoculars (ls. 972-977) Crotais foonacus [No transverse bar in the prefrontal areas as described 2 23, Toil usually Black or very dark brown or gray with pale crossbars rarely in evidence on posterior of tall rate matrix usually black (Pls 876-888) Crotalus molossus* Tail usually gray with pale gray crossbars in evidence posteriorly; rattle matrix usually gray or brown (Pls 781-787) aa Crotalus baslscus 4. Minimum number of intersupraoculars 4 or fewer (PL. BAD) ...e00 (Crtals lannon Minimum numberof intersupraoclars 5 or more (PI. 967) seoevee Clas steegeri 1. This ys dap npr to Gly 190 and evel ays pci in Kay, 1972 Whar ed any of he sac eae a Ler nes by lye 5} a Rue C9 foe etn hey orate The Cerra ae ear ata be er aml elon qresiga segltoy Ue al adel ‘Sher ning sie renal eter and westin Unie St, eo the Meso ops rs n mone Naar de pen erate eats’ nt deseone al many morpholopal its fave en Sas wor comely pcos cen ci fe ata Vary whether ott eine sca oath april ser. aia spas Cie pes and Cea a ay nas Sater We fave fone ber C973), Seneca ets ecole C pic Gp pecinen ows orn vw te air uel a sSreach Resi Me etna oi Trias mmr hare prvi tc ragtne i neotenic of. e. Campbell tt Estar (1983) wa the frst to alloca these poptlatons te € nofasus pune, Previous Confusion of Oaxacan tepatmn wit ftecre py sae at enn pr ese Omen nas ed ohsve easy tt Bi gec fyens oc vergaemnotercali yar wala enypnecy Clave para las Especies de Serpientes de Cascabel! 1. Dorso de Ia cabeza con menos de 12 placas grandes, distribuidas simétricamente & incluyendo internasales pares, prefrontales pares, y una frontal tnica : 2 DDorso de la cabeza con mas de 12 placas de menor tamafo, distribuidas asimétricamente e incluyendo varias fen la rogicn frontal y nuimerosas escamas en la regin parietal seco 2, Canto angularcon ola lateral deescama canta con un filo moderado, no curva hacia abajo al lado dela cabeza; rostral alla, truncado de ariba; manchas corporates usualmente tan anchas 0 mas anches que largas 2. Canto redondeado con orils lateral de eseama canta curva hacia abajo al lado de la eabera; rostral baja, pune feando al curvar posteriormente entre las internasales; manchas corporales uswalmente més largas que anchas (Ps. 931-839) ares fish Crotalus raous 3, Prevcular superior usualmente en contacto con postnasal, subocular anterior usualmente contacta supralabi- ales 4 y 5 (Pls 1011-1017) 3 Sisturus cutenatus Preocular superior usualmente no en contacto eon postnasal, subocular anterior usualmente contacta supral- ablales 3 y 4 (Pls. 1018-1023) evseoceneesse Stuns mins 4. Orillaslaterales de supraoculares elevaclasformando procesos planos, puntiagudas y en forma de enero (Ps 791-794) ‘Cotas corastes Orillas laterales de supraoculanes no extendiéndose para formar procesos puntiagudes yen forma de cuemo sees ese ; 5 5. Escamas subcatdales en machos menos de 40 y en hembras menos dle 35, 6 Fscamas subsaudales en machos mas de 40 y en hembtas mas de 33 : au 6, Punta del hocico y porcin anterior del canto clevado formando un filo agudo; rostral y mental muchas veces com una linea vertical media palida (Pls. 998-1010) + Crotalus zl ‘Panta del hocico y porcin anterior del canto no elevado formando un filo agudo; rostral y mental sin una linea vertical media pala ne 7, Pronasales separadas de rostral por pesjueias escamas, muchas veces granulares;preoculares superiores muchas ‘veces dividicas horizontalmente, verticalmente, o de ambas formas (Pls. 864-875) Crotalis mith Rattlesnakes 487488 10, 2. 1B 18, 6, 2. 2. Prenasales contactando rostral: preoculares superiores usualmente no divididas 0, si asf es, dividicas dinica- mente verticalmente con la porcion anterior de la escama consplcuamente ms alta que la posterior y curva sobre el canto al frente de a supraocular «... 5 Una serie de 35 0 mis bandas transversales formadas de puntos oscuros en un fondo mis palido, ancho dorsoventral del segmento proximal del cascabel menos de 25 veees en ol lago de Ia cabeza (Pls. 968 y 969) . Crotalsgrs Manchas corpoales dorsales en forma de diamantes, hexigonos, ectingulos, vals, 0 elipses 0, si con bandas teansversales presente, estas no formadas por puntos conspicuos, ancho dorsoventral de! segmento proximal dda cascabel mis de 2.5 veces en el largo de la cabeza Subocular anterior en contacto con To mis (usualmente 2) supralabiaes 10 ‘Subocular anterior no en contacto con cualquier supralabial v7 Supraoculares atravesadas tranaversalmente por una linea pala y delgada bordeada de negro; con una mancha redonda u ovalada bajo el ojo que esti delineada distintivamente; patron dorsal corporal de elipseslongitudi- rales (Ps. 915-918) panes ‘Crotalus plysictus Suptaoculares no atravesadas transversalmente por una linea plida y delgada bordeada de negro; sin una ‘mancha reionda u ovalada bajo el ojo que esté delineada distintivamente; patron dorsal corporal no de elipses Tongitdinales 11 Hileras de escamas 3 mitad del cuerpo més supralabiales en ambos lacos suman 41 6 menos 2 Hileras de escamas a mitad del cuerpo més supralabiales en ambos lados suman 42.0 més 4 Loreal en contacto can 1 0 mas supralabiales (Pls. $35-838) Crotalus intermedius Loreal no en contacto con cualquier supralabial 1B PPreocular inferior pasa ala fosa facial y no logra hacer contacto (o solamente de forma muy angosta) com la loreal; patron corporal de bandas transversas dorsales no divididas Cotas transcersis Preocular inferior pasa arriba de la fosa facial y hace contacto con la loreal; patrén corporal usualmente de ‘manchas apareadas (algunas de las cuales pueden estar conectadas ireegularmente através de Ia linea media dorsal (Ps. 919-928) Crotalus price’ Solamente 2 escamas en el rea prefrontal (Pl. 929 y 930) Crotalus pusilus Mas de 2 escamas en el rea prefrontal : fas Proocular superior usualmente dividida verticalmente, la seecin anterior siendo mas alta que la posterior y curva sobre el canto en frente de la supraocular; manchas dorsales corporales (menos de 25) ccupan menos ‘espacio longitudinal que los espacios entre llas (Pls. 841-863) Crotalus leis Preoeular superior io dividida verticalmente 0, si dividida, Ja seccién anterior no siendo mas alta que la posterior y no curva sobre el canto en frente de la supraccular; manchas dorsales coxporales (25 0 més) ‘ocupan mis expaci longitudinal que los espacios entre cllas .-.ee-e+o.- 16 ‘Usualmente 10 escamas bordeando el cascabel; cascabel relaivamente grande, en serpientes excediendo los 40cm de largo total, el ancho dorsoventeal del segmento proximal del cascabel es mas de 1% del largo total del cuerpo (PIs. 756-772) Crotalus aquilus LUsualmente § escamas bordeando e] cascabel: eascabel relativamente pequefio, en serpientes excediendo los 440m de largo total, el ancho dorsoventral del segmento proximal dol cascabel es menos de 1%s de largo total del cuerpo (PIs. 981-987) : ae Crotalus tiserits ‘Mis de 2 escamas ente las nasales en contacto con rostral (usualmente nombradas internasales) (Pls. 889-914, 988-997) covsesee (Crotalus oreganns, . viridis (ver descripciones) ‘Dos escamas fintemasales) entre nasales en contacto con rostral eseeree8 Preocilar superior ustalmente dividida verticalmente la seecién anterior siendo més alta que Ia posterior y ‘curva sobre et canto en frente de la supraocular; manchas dorsalescorporales ocupando menos espacio longi- tusdinal sobre la linea media que los espacios entre elas (Pls. 1-863) Crotalus lps Proocular superior no dividida verticalmente 9, sidividida, la secei6n anterior no siendo mas alta que la posterior y no curva sobre ol canto en frente de la supraocular; manchas dorsales corporales ccupando ms ‘espacio longitudinal sobre la lines media que los espacios entre elas 19 ‘Ventrales 163.0 menos a 2 Ventrales 164 0 mis : a Usualmente 10 escamas bordeando el cascabel; eascabel relativamente grande, en serpientes excediend los ddtem de ltgo total el ancho dorsoventral del segmento proximal del cascabel es mas de 1% del largo total det ‘enerpo (Pls. 756-772) Crabs agus ‘Usualmente 8 escamas borleando el cascabel;cascabel relativamente paqueo, en sexpientes excediendo los 440m de largo total, et ancho dorsoventral del segmento proximal del cascabel es menos de 1% del largo total ‘dol cuerpo (Ps. 981-957), Cros trsriats Bandas o anillos de la cola fuertemente contrastantes y conspicuos, blanco o grs-blanco altemacio con castaio ORCUTO O HERO 2 Cola negra o gris oscura, bandas o arilles, si presentes, solo moderadamente evidentes debido a ta falta de ccntraste entre bandas o aillos oscutos y el fondo 2 Pitvipers, Viperidae122, Matriz del cascabel encogida, un solo segmento de cascabel, sin segmentos sueltos (PIs, 788 790). eae (Chota calainensis Matriz del cascabel normal, usualmente varios segmentos de cascabel presentes (excepto en juveriles muy ppequerios que pussen tener solamente un bot6n) . B 23. Anillos o bandas oxcuros de la cola como de igual o mayor longitud que los espacios palidos entre ellos; mimero {ninimo de excamas entre las supraoculares raramente menos de 4;sin escama plana en forma de lina ceciente bordeanda posteriormente cada supraccular 4 ‘Anil oscuros de la cola de menor longitud que los espacios plidos entre ellos; timero minimo de escatmas Gitte [ss supraceslares raramente usualmente 2; usvalmente con 1 escama plana en forma de luna creciente bondeando posteromedialmente cada supraocular (PIs, 945-955) Crotalus seutatas 24 Una barra pélida vertical en prenasal. y_ primera supralabial; linea postocular pélida, extendigneose Gente el bore posterior del ojo liagonalmente hasta tun punto arriba o deirés del éngulo de la boca (Ps. 732-753) ‘Crotalus adamarteus ‘in barra palida vertical en prenasal y primera supralabial linea postocularpalida exten dignse desde el bode posterior del jo diagonalmente hacia abajo hasta ultima o penitima supralabial 25 25, Primer par de infralabiales usualmente divididas transversalmente; manchas dorsales no conspicuamente mareadas con puntos negros e 26 Primer par de infralabiales usualmente no divididas transversalmente; manchas dorsales conspicuamente ‘mateadss con puntos oseures : . 7 26, Uswalmente sin escamas intergeniales; prenasal usualmente contactanclo primera supralabial;anillos escuros de a cola usualmente no intersumpidos lateralmente (Ps, 40, 941, 945-948) ‘Crotalus ruber UUsunlmente con escamas intergeniales; prenasal usvalmente no en contacto con primera supralabialy anillos ‘curds dela cola usualmente inerrumpidos lateralmente (fsa Cedros; Pls. 62-948) Crotalus ruber 27, Preocular superior usualmente en contacto con postnasal 0 ese contacto impedico por una loreal superior (PIs 773-783) as zs Croats atrox Preocular superior usualmente no en contacto con postrasal y sin loreal superior presente (Pls. 970 y vr) (Crotalus tortugensis 28, Patedn corporal de bandas transwersales negras en forma de V (Pls, 829-834)... Crotalus horrus Patrd eorporal de manchas en forma de diamantes, cuadros, 0 edondeados 29 29, Regis internasal-preftqnal de la cabeza cubierta con 13-25 escamas; corolla y ea frontal eubiertas mayor mente dle escamas éeperas,quilladas, y protuberantes (Pls. 825-825) Crotalus eryo Regn intemasal-prefronta'de la cabeza wsualmente cuberta de 12.0 menos escamas; cronillay érea frontal ccubiertas mayormente de escamnas planas y lisas 0 ‘Con un par de rayas dors laterales destacaclas en el cucllo de 1-3 escamas de ancho, separadas pot un espacio pilidontedio dorsal de 2-3 escamasde acho, y extendiéndose I argos de a cabeza ders de a cabera antes de llegar a a primera mancha dorsal : 31 Fr Sin un par de rayas oscuras en el cuello 0, si presente, iregulares y extendiéndose posteriormente menos de tun lange de cabeza detrés de la cabeza : 2 431, La mayoria de las manchas dorsales en forma de diamante separadas lo largo de la linea media dorsal y tha extrnditindose hacia abajo por los lados como bandas transversales; lineas paravertebrales en el cuello Pondeadas lateralmente por una hilera de escamas pélidas que no son conspicuamente mas palidas que el Srea adyacente inferior (Ps. 956-968) (Cras sims Manchas dorsales en forma de diamante mayermente en contacto a lo largo dea nea media dorsal lineas par~ herebrales en el euclo bordeadas lateralmente por una hilera de escamas plidas que son conspicuasmente Inds pilidas gue ls de Ia hilera adyacente inferior a los lados del cuerpo (PIs. 798-824) ..... Cots durssus 132, Una barra negra o castalo oscura con borde palo eruza la cabeza entre los puntos anteriores de las supra ceulares (Pls. 972-977) Crofals totomacus Sin barra transversal en las reas prefrontales como descrta anteriormente 3 33, Cola ustalmente negra o eastaiio muy oscuro o gris con barra transversal pilidas raramente evidentes en [parte posterior de la cola; matrz del cascabel usualmente negra (PIs, 876-888) ‘Cras moss? Cola usualmente gris con barras transversales gris plidas evidentes en parte posterior; matriz del cascabel tusualmente de color gris 0 eastaio (PIs. 784-787) : : Crotalus tsilseus 34. Naimero miniino de intersupraoculares 4.0 menos (Pl. 840), (Crotels lnam [Naimero minimo de intersupraoculares 5.0 mis (Pl. 967) ‘i Crotalus stejneger 4, Tata dave es adaptada on parte de Gloyd, 1940 y varias lave presentadas en Klauber, 1972 Hemos usado muchas de lad niche sataceritket externes que fueron usadas por Gloyd (1940) y Klauber (1972) en sus claves de identificain para eile de cate La clave presenta act sn embargo cifire de las aerores en varios aspects significative una aries (lupe todas ins serphrtes de casa en lugar de estar subividias regionalmente (ese y dete de ls Estados Tee dieah i eopecte Mexcana reo es movida a Cott; nuevas especies de Crotls son teconxidas: a variacin J aandcon de muchos caavteres morfolegios han sido pusts al dia sag lo permite el conicmiento Saat cupeles de sepients de cvtabel pegs sn variables con respecto a ener ono la subocular anterior conta ee ettesapraabotes. Coteus aj, eps, Cpa, y C sera son wanables con respect # este cara Rattlesnakes 489ter Hemos seguidoa Klauber (172) en presenta doblementea estas especies con excepcion de C. psi em os que un 00 ‘jemplar es cnocido porno tener Ia sbocular anteiox sin yar las supralabiles “T'Poblaciones surenas de Crviies wlesus han sido. previamente reconocidas como subspecies alopitricas de C Isls Campbell y Lamar (1983) faeron los primeros en Klentfcar estas poblaciones convo Cais moss aaacus, Con sn previa di las pobliclones de Oaxaca con C. Boss probablemente surgi, al mencs en pare, porque ls sempientes dle Omuca tienen a fener anos catdaks relativamente destacados; este earacter aparentemente vara un poco clial y ‘ntogeseticamente Rattlesnakes, Genus Crotalus Linnaeus, 1758 Choialis Linnaeus, 1758, Syst: Nn, 10h ed, 828 p21. Type- ‘species: Crotalus lorridvs Linnaeus, 1758, by subsequent

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