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English Pages [576] Year 2016
Peter Heimes
Herpetofauna Mexicana Vol. 1
Snakes of Mexico
FRANKFURTER BEITR.AGE ZUR NATURKUNDE BAND70 FRANKFURT CONTRIBUTIONS TO NATURAL HISTORY VowME70
Peter Heimes
Herpetofauna Mexicana Vol.
1
Snakes of Mexico
Edition Chimaira c!_�� Frankfurt am Main �
Pecer Heimes Herpetofauna Mex.icana Vol. r Snakes of Mexico Edition Chimaira Frankfurt am Main 2016 ISSN 1613-2327 ISBN 978-3-3 899 73-100-2
To my wife Sylvia: This is your book, because without you none of this would have happened.
On the front cover, top: Leprophis ,1haewila, Caremaco, Sierra de Los Tuxrlas, Veracruz, photo by I� HEIMES; middle left: Crotalus sc1tt11!ruus snlvini, El Frijol Colorado, Perore, Veracruz, photo by P. HEIMES; midclle cenue: Lamprope!tis knob!ochi, Sierra Tarahumara, Chihuahua. phorn by P. HEIMES; midclle right: Adelphicos 11igrilat11s, Mit2ir6n, Chiapas, phoro by C. I. GR0N\VAI.D & J .M. ]ONES; bottom left: Lamprope!tis calijomiae, Ensenada, Ba ja California, photo by P. HEIMES; bottom right: Lichanura r:rivirgata trivirgara, Jose del Caba, Ba ja California Sur,
photo by M. CAGE. Design by Gerold SCHIPPER.
All rights reserved. No part of this publication may be reproduced. scored in a recrieval S)'Scem or cransmicred, in an)' form or by any means, elccrronic, mechanical phococopying, recording or otherwise, without the written permission of che copyright holders.
© 2016 Chima.ira Buchhandelsgesellschaft mbH Publisher: Andreas S. Brahm Heddernheimer LandsrraEe 20 60439 Frankfurt am Main Germany Phone: +49-69-497223 F ax: +49-69-497826 E-Mail: [email protected] Website: www.chimaira.de Layout: Balazs Farkas, Gyur6, Hungary Priming: Colordruck Solutions GmbH, Leimen, Germany Binding: Freitag, Kassel, Germany fSSN 1613-237-7 ISBN 978-3-38997:>-100-2
In the US available from: ECO Publishing Bob and Sheri Ashley 4 Ratclesnak.e Canyon Road Rodeo, NM 88 056 Phone: (575)-557-5757 Fax: (575h57-7575 E-mail: [email protected] Website: www.ecouniverse.com
Contents Foreword by Larry David W1LSON .. . .. • ...... • ... . • . • . • .. Introduction . . ... ..... ....... ............... . ....... Acknowledgements ....... , .......... ................. The Natural Environmem of Mexico ..................... Composition and Pattern of Distribution of che Mex.ican Snake Fauna ...................... ,..................... Species Accounts .......... ....... ......... ...... ..... Infraorder Scolecophidia .......... , .. ........ ........ Family Lepmryphlopidae ... ......... .... ..... ...... Genus Epictia ... .... ........... • ..... • ... • . • . .. Epictia goudotii . ...................... ... ..... Genus Rena ......... . ......................... Rena boettgeri ....... ..... • .......... .... ..... Rena bressoni .... • .... ..... .. • .... .... ...... .. Rena dissecta ......... ........................ Rena dulcis ........................ , . , ... , . .. Rena humilis ..... ........... . .... ... ....... .. Rena maximt1 ..... • ... • ... • . • ...... . . • .. • .... Rena mJ1opica ................................ Rena segrega ..... . . ......... ................. Family Typhlopidae ... ......... ......... ...... .... Genus Ameroryphlops . • ....... ............... .... Ameroryphlops microstomus ............ · . ......... Ameroryph!ops tenuis ...... , .......... .......... Genus lndm)'phlops . ....... ...................... lndoryphlops bramimis . ... ................. • . • .. lnfraorder Alerhinophidia .. ......... .... ............. Family Boidae ......... .......................... Genus Boa ....... .... ..........•..... • ........ Boa constrictor ... .•......... .. ..•. • • .. . ... . .... Fa111ily Charinidae . . .... ..... . .......... . • ........ Genus E·dliboa .....•.............. , .. . • .....•.. Exiliboa placata ........ .... ... ......... ....... Genus Lichammt .. ... .... • . ..... • ... . . • .. • . . • .. Lichmmm trivirgata ......................... .. Genus Ungaliophis ....................... , ... . .. Ungaliophis co11tinentalis .............. ...... ... . Family Loxocemidae .... ... ..... ................. . Genus Loxocernus ......... • ....... .... ......... . loxocemus bicowr ................... .......... Family Colubridae ...... ..... ....... .... . .... ..... Genus Arizona ........... • . • ..... .............. Arizona eLegans .......... ..... . .. . .. . .. . ...... Arizona pacata .... . ......... . • .... . ...... .... Genus Bogertophis ............... , ............... Bogertophis rosaLiae .. .......................... Bogertophinubocularis ..... . • ... • ............. .. Genus ChiLomeniscus .................... ...... .. ChiLome11iscus savagei ...... , ........... ........ Chiwmeniscus stramineus ........................ Genus Chionactis .... ... ... ..... • .... ........... Chionactis occipitaLis ... ........................ Chionactis palarostris ............... . ......... .. Genus Coluber ...... ... .... . ................... Coluber constrictor .................•............ Genus Conopsis . ... , ............................ Conopsis aettta ........ .... • ........ ......... .. Conopsis amphisticha ... . • .. • ....... ............ Conopsis biseriaLis .............. ............... Conopsis lineata .................... ........... Conopsis megalodon . ... ..... ....... ...... .... .. Conopsis 11ams ......... , .. , .. , .. • ............. Genus Dendrophidion ............................
9 10 12. 13 19 11 26 26 26 26 27 2.7 27 29 29 30 30 31 31 34 34 34 34 35 35 37 37 J7 37 39 39 39 39 42. 42 42. 45 45 45 47 47 47 50 50 50 51 51 54 54 55 55 55 58 58 59 59 62 61 62 63 63 65
Dendrophidion vinitor .......•......... , .... ... . 65 Genus D rymarchou ..................... , .... .... 65 Drymarchon melnnurus .... • ..... • ... ........... 67 Genus Drymobiw ............... ............ .... 68 Drymobius chLoroticur ....... . • ........ • ..... ... 68 Drymobius margaritiferus .... ................... 69 Genus Ficimia .. .. ....................•.. ...... 69 Ficimia hardyi .......................•.. , . .... 72 Ficimia olivacea ..... ................ • ........ 72. Ficimia publia .................. .............. 72. Ficimia mmirezi ..................... ... ...... 73 Ficimia mspator . ............... ............. . 73 Ficimia streckeri ... .... ......... . . • ........... 73 Picimia variegara ...... .. • .... ................ . 76 Genus Ceagms .. ......... • .... ................. 76 Geagra.s redimittts ....... • ..................... 76 Genus Gyalopion ......... • ........ . ..... . ...... 77 Gyalopion canum ......... ... .......... ... .... 77 Gyawpion quadrangulare ............. ....... .. .. 79 Genus Lampropeltis ............................. 79 Lampropeltis abnorma ................•. • ....... 80 Lampropeltis altemri ........................... 81 Lampropeltis mmulata .......................... 84 Lampropeltis califomiae ....... .............. . ... 84 Lampropeltis catalinensis . . . . • ..... . .... . . . .. • ... 85 Lampropeltis knoblochi ....... .................. 85 Lampropeltis mexicana .. ..... ................ .. 88 LampropeLtis poryzona ........ ............. . • . .. 89 Lampropeltis ruthveni .................. • .... ... 89 Lampropeltis splendida ........ .................. 92. Lampropeltis webbi .......... ... ............... 92 Lampropeltis zonara ................ ......... ... 93 Genus leptophir ..... . ....... ....... . . .......... 93 Leptophis ahaetttlla ....... .. ........... • .... ... 9 6 Leptophis diplotropis ........................... 96 Leptophis mexicamts ........ ................ ... 98 Leptophis modestus .... ... . • .... .. .......... .... 99 Genus Masticophis ............. ................. 99 Masticophis anthonyi .. ......... ................ 102. Masticophis aurigulus ........ ... • ..... ......... 102 Masticophis barbouri ............ , ............ . . 103 Masticophis bilineatus .......................... 103 Masticophis flagellum ..... ........... .. ... ... ... 105 Mrtsticophis fitligi1wms ........................ , . 106 Masticophis latera/is ............. • .............. w7 Masticophis mentovariw ........... ....... ...... 107 Masticophis schotti ................... .......... 110 Masticophis s/evini .................. ..... ...... 111 Masticophis taeniatm ................... .. .... • . m Genus Mastigodryas ..... • . • ........... ....... ... u4 Mastigodryas cLiftoni ........................... t14 Mastigodryas me/anolomus ... .............. .. .... 115 Genus Opheodrys ..................... . • ..... .. . n6 Opheod1ys aestivm ......... ................ .... 116 Opheod,ys vemalis ...... .................... .. . 1.16 Genus Oxybelis ............................. .... 117 Oxybelis aeneus ..... .... . ............. ... ..... 118 O>.ybelis fitlgidus ..................... • .... .... 119 Genus Pantherophis ............ ........ • ....... . Tl9 Pantherophis bairdi ...... • . .... • .......... . .... 112. Pantherophis emoryi ..........•.•...... , .... .... 12.2 Genus Phrynonax ........... ................ .... 123 l'hrynonax poecilonorus ... • ................. .... 123
Contents
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Genus Phy!Lorhynclms ............ , ... ....•....... Phyllorhynchw broumi .......... • ... . . . ......... Phyllorhynchus decurtatus ........ . • ............. Genus Pitttophis .. ..... . ........................ Pituophis cateuifer ... . .... ...•................ . Pituophis deppei ................ . ...... ...... . Pituophis lineaticol!is .................... ....... Pituophis vertebra/is .... ........................ Genus Pseudelnphe .. ...................... ...... Pseude!aphe J!avimfa ............... ... . . ...... . Pseudelnphe phnescens ........ . . .... ..... ....... Genus Pseudoficimia .. .. .. . ...•.................. Pseudoficimin frontnLis ............. . ............ Genus RhinocheiLus .................. .. ........•.. RhinocheiLus etheridgei .............. .........•.. RhinocheiLus lecontei . . ................ •........ Genus S11lv11dora ....................... • ...•.... S11Lv11dora bnirdii ............ . . ..... • ... ....... S11Lv11dom deserticola ......•.... .... • •.......... Salvadom gmhamine ............•...•.......... Salvndora hexalepis ... .......... • ....... ....... SaLvndora intermedin .. .......... . ....... ....... S11!1111dom lemnismtn ...•....•......•.... • .. • ... SaLvndora mexicana ........ ............. •...... Genus SenticoLis .... ........ . ................... Senticolis trinspis .... • . ............. ........... Genus S011ora ...... . ................. • ...• ..... Sonom nemu!a. .. . ..... .................. ..... Sonom michoacanensis .......................... Sonom mutnbilis ......... .... ................. Sonom semi11n111t!ata . ......... ........ .. . ...... Genus Spilotes ..... ... .. • ... . ................ ... SpiLotes puLlattts .. .......................... ... Genus Stenon-hi1111 ........................•..... Steno,-,.hi11a degenhardrii ......... ............. . . Stenon-hina f-eminviLlei .................... . ... . Genus Symphimus. ..............•...•.•.•....... Symphimus leucostomus ......... • .•...•...•..... Symphimus mayne .............. .•..... . .. • ... . Genus Sympholi.s ............ • . • ...•.....•....... SymphoLis lippiem . . ..... ...................... Genus TantiL"1 ................................. Tantilla atriceps . .................... . ..... .... Ttmtilla boco1mi ...... ... ...... ... ........ .... Tantillo briggsi ......... . .. .... ........... .... TantiLla calamarina ............................ Tantilla casmdae ..... • ...... ..... ........... .. Tantilla ceboruca ............... • .• . .......... . Tanti!!a coronadoi .. • .• • ..•...• ... ............. Ta11til/a crmiett!ator ............ ....... . ........ Tantil!a deppei ................ .•............ .. Tamil/a flaviLi11eata ...... ... • . . .......... . ... . . Ttmtilla gracilis ............•.....•. .......... . Ttmti!la hobartsmithi ......... . ..........•.... .. Tantillo impema .............................. Ttmtilla Jani ....... ...... .......... • .....• . .. Tant·illa johnsoni . ... .... . . ....•. ....•......... Tnntilla moesta .. ............................. Tantilla nigriceps ............................ . . Tantillo oaxame. •..... ............ • ...... .. ... Tantilla planiceps ........•............... ...... Tantilla robusta .............. .............•... Tantillo. rubra .................... , .•. • .. . .... Tantilla schistosn .................... .......... Tantilla sertula . .... . .... .... . . ............... Tantil!a shawi ........ ........................ Ta.ntil/a slavensi ...............................
126 126
127 127 130 131
134 134 135 135 138
138 138 139 139 142
143
143
146 146 147
150
150 151 151 151
154 1 55
155 158 1 58
159
162 162
162 163 166
166
167 167
167 169 170
171
171 174 174 175 175 1.75
178 178 178 179 179
181
18r 182 182
183 183 183 184
184 185
185 187
Tantilla striata ......................... ...... 187 Ta11tilla tayme ......... ....................... 18 7 Tamil/a triseriata ...........•...... , .. ... ...... 188 Tamil/a 1111/cani ......... ......... .... • . ...... . 188 Tantilla wilcoxi ........ . ...... .. . ............. 189 Trmtil!a ynqt,ia .. . • . .......................... 189 Genus Tantillita .. ................ .............. 189 Tantil!ita brevissima .................. . ........ 191 Timtillita camt!a ............. .......... ....... 192 Ta.ntif!itn lintoni . ........ ............. ........ 192 Genus 1i·imorphodon ... . ............ ....•.... ... 192 1i·imorphodon biscutatus .................... .... 193 Trimorphodon lambda ............. • . ..... . ..... 196 1i-imorphodon lyrophmies . ............ ... ........ 196 1i·imorphodon paucimaculatus ......•...... ....... 197 1i-imorphodon tnu ....... ......... ...... ....... 197 Trimorphodon vilkinso11ii .................•...... 200
Family Dipsadidae
, ........ ......... ......... ..... 101 Genus Adeiphicos ........................ , .•.. .. 201 Adelphicos latifasci11t11s .... .............. ...... .. 101 Adelphicos nigrilatus .......... .......... . . .•... 204 Adelphicos quadrivirgatm ... • . .•....... .... ..... 204 Adelphicos sargii ............................. . 204 Adelphicos visoninus .. ..... . ...•.•........ ..... 205 Genus Amast1·idi11m ........ ....... • ............. 205 Amnstridium snpperi .... . .... . • . .•. ............ 205 Genus Chersodromus . ..................•.•....... 208 Chersodromus liebma1111i ........................ 208 Chersodromw rubriventris .............. .•....... 208 Genus Cleli11 . ..........•....................... 209 Clelia SC)'trdina ... ......... ......... .......... 209 Genus Conioph111w ........•.•.................. 209 Coniopbrmes 11lv11rezi ........ ................ ... 212 Co11iophanes bip,mctaius .. .... • ..• .............. 212 Co11ioph1112es fissidens ......•................•... 213 Co11ioph1111es imperialis . .........•. . ........... . 216 Co11iophanes lateritius ............. ............. 216 Co11iophanes melanocephflius ..... .•... ... ........ 217 Co11ioph1111es meridanus .............. ........... 217 Coniophanes michoac1111ensis ..................... 217 Coniophanes piceivittis ................ . ......... 220 Conioph�nes qui11que11ittatus . .......•. ........... 220 Co11iophm1es same ..... .... .................... 221 Coniophanes schmidti .. • ....................... 221 Coniophanes ta)'lori . . ... ............... ........ 221 Genus Conophis ....................•....•...... 224 Conophis linentm •................. .. ......... 224 Conophis vittatus ....•... • .... ...... .. . ........ 225 Genus Cryophis ............ ....... . . .... •...... 225 Cryophis hn/Jbergi • .. • . . • ... • . • • . • • . • • .... . . . .. 215 Genus Diadophis ...... .... .•... • ............... 227 Diadophis p1111ctat11s .. .......•........ ......... 227 Genus Dipsas . ........ ........... •..........•.. 128 Dipsa.s brevifacies ..... .............•...•. • ..... 228 Dipsas gaigeae . ...................... .•....... 229 Genus Enulius ........•.......... . . ............ 229 Emt!ius flavitorques ............................ 229 Enulius oligostichus ...... . .. ...... ............. 231 Genus Geophis .... .... .................... ..... 232 Geophis anoculnris . ....... •... . .•.............. 233 Geophis bicolor ..............•.•.. , .....••.... 233 Geophis blanchmdi ....•.... ..•...... ... ...... . 236 Geophis mncellatus .........•.... .....•........ 236 Geophis carinosus .... ... .................... ... 236 Geophis chalybeus ...... ....................... 236 Geophis dubius ........... - . - - .. - ..... ........ 237 Geophis duellmani ......................•...... 237
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Concencs
Geophis dugesii ........•........... ... ........ 237 Geophis immnml11tt1S ....•.......... ....•.....•. 240 Geophis incomptus ...... • ... .... .. ... ... ....... 240 Geophis isthmirns ...... ..• ... ... ............... 2.41 Geophis juarezi .. .......... • ......... ......... 241 Geophis juliai ...... ........ ... . . .... ...... ... 241 Geophis laticincttlS .... ......................... 241 Geophis !aticollaris ....... ... ......•........ .... 244 Geophis !atiji-ontalis ...........•. , . , ............ 244 Geophis maculifarus ......... ....•..... • ........ 245 Ceophis mtttitorques ..... .....•........•... .... . 24 5 Geophis nnsnlis . .. . .... .................... . .. 245 Geophis nigrocinctus .................. . . ........ 248 Geophis occabus ...... , . .•.. ............. ...... 248 Geophis omiltemarms .... . • . ............... . .... 248 Geophis petmii .........•... ............. ..... 249 Geophis pybumi .............. .•....... ........ 249 Geophis rhodognster .. . ........ .. ...........•... 249 Geophis rostmlis ............................... 250 Geophis sallei ............ .....•.... ........... 250 Geophis semidoliatus ...... .........•. • ......... 250 Geophis sieboldi .. , .. , .. , .. ..•.......•.... ..... 25-r Geophis tnmscae. ......... ...... ......... ...... 25'1 Geophis t11rbidus ... ............ .•...•......... 251 Genus Heterodon ........ .. ...... ....... .. ...... 252 Heterodo11 kennerlyi ... .. • ...................... 252 Genus Hypsigle11a .......... ............... ...... 252 Hypsigle11a affinis • ................. ....• ....... 153 Hypsiglena catd{inne ..................... ... • . •. 253 Hypsiglena chlorophaM . , .••...... , •.... , •. , .. , . 256 Hypsiglena jn11i ......... ...... .... ..... . ...... 256 Hypsigleua ochmrhyncha .. . ......... . ..... ...... 257 Hypsigle11a slevini ........ , .................... 260 Hypsiglena tanzeri .. ............... .. ...• ...... 260 Hypsiglena torquam .......... , .... , . . .... ...... 260 Hypsiglent1 11/ltlocult1ru.s ........ • ... ............. . 261 Genus lma11todes .................. , ..... ....... 261 lmantodes cenchoa ........... , •. , .. , ..•.. , ..•.. 264 fmantodes gemmisrratus ................ • . .... • .. 264 lmamodes te11uissi111us ................. , . ....... 265 Genus Leptodeim ........ ...... ... ... . .. . .. ..•.. 265 leptodeim ji-enata ....... ............... .... . .. 268 leptodeira mamlata .... ............... . ....... 269 leptodeira nigrofascit1ta .................. .... ... 269 Leptodeim polysticta .. ... ...... ......... ........ 272 Leptodeim punctata , .. , ........................ 272 Leptodeira septenn·ionalis . ...... .......... ....... 273 Leptodeira spleudida ........................... 273 Leptodeim uribei ........ , .. , . ......... ........ 276 Genus Manolepis .........•..................... 177 Manolepis putnami ......•.................•..• 277 Genus Ninia.. ...........•..•.. , ............... 277 Ninia diademata ............•.•........... , .. , 280 Niuia sebae .• , .. , .. , ......... , ........... , ... 280 Genus Oxyrhoptts ............ , ..•..•............ 281 Oxyrhop11s petola ... ...................... ..... 281 Genus Pliocerrns . , .................... , ..... , .. , 283 Pliocercus elapoides . ....... . ..... • ... . .... ..... 283 Genus Pseudoleptodeira ......... ........ •...•. •. .. 284 Pseudolcptodeira larifasciata ..... ......... ........ 285 Genus Rhndinaea ..... , .............. , , ... ...... 285 Rhadi1111ea bogertorum , .. , .......... ..•......... 288 Rhadi1111e11 cttneata ........•..... , . .. • ....... .. 289 Rhadinaea decomta ......................... ... 289 Rhadinaea forbesi .................. ..... .. • ... 292 Rhndinaea Jitlvivittis .... , .. , . , , ........ , .. , .... 292 Rhadi11aea gaigeae ............ ..... ..... . ....• . 293
Rhadinaea hespcria ....................... .• ... Rhadinaea laureata ........ . .......... ...... .. . Rhadinaea macd()ltgalli ..... , .............. ..... Rhadinaea marcellae .... , .. , ................... Rhadinaea montana. .......................... , Rhadinaea myersi . ............... • ............ Rhadi1111ea. omiltemana .......... .... . ...... .... Rhadinaea quinquelineata .. ...... ............... Rhadinaea taeniata .... ......... ............... Genus Rhadinella ........•...................... Rhadinella donaji ..•..... , ..... .. ......... .... Rhadinella godmani ... ......•... •.•....... ..... Rhadinel/11 hannstei11i .. ......... • .•............ Rh11di11ella kanalchutchan ........ • . ... .... ...... Rhadinel!a kinkelini ..............• ...•........ Rhadi11ella lachrymans .........•.. .....•........ Rhadinella posadasi ................. •. • ........ Rhadinella schistosa. ............ .. ....... ....... Genus Rhadinophanes ............................ Rhadinophanes monticola................... ..... Genus Sibon .... .. ....... ............ ..... • .... Sibon dimidiatus ..... , .. , ..................... Sibo1z linearis . . ..•...•....................... Sibon nebulatt1S .......•.•.....•.... ........... Sibon sannio/11 ..... • •.........•.............. . Genus Tantalophis ............. .. .............. . Ta11t11lophis discolor ..................... , ...... Genus Tretanorhinus .. ............... •.... ..... . . 7i·eta11orhi11w nigroluteus ... . ........ ....... .... . Genus Tropidodipsas ...................•..... , ... Tropidodipsas mmulifara ......... ............... Tropidodipsasfasdattl ............•.. ........... Tropidodipsas fischeri ................ ... . ....... Tropidodipsas philippii . ........ , .. , .. • ..• ... ..... T,·opidodipsas repleta. ........................... Tropidodipsas sartorii .......... , ..... , ...... , ... 7i·opidodipsas zweifali ... ....................... Genus Xenodon ...................... .... . •.. . . . Xcnodon rabdocephalus ................ ... ... ...
Family Elapidae .......................... ... .. ... Genus Micruroides ...... ........ .... ...... • ... .. Micmroides ewyxamhus ............. , . , .. •..... Genus Micmrus ..•. .... . .... .............. ..... Micrurus bemadi ..•......................•. . . Micmrns bogerti .........•........... , ........ /vficrurus browni .............•...• , •. , ........ Micrums diastema ............ ................. Micrums distam ....... ..•.................... Micrunes elegam .......•.•........... , ......•. MicrtmtS ephippifar .......•...•.•..... , .... .... Micru,w laticollaris ................... , ........ Micru,w !atifosciatus .................. , ..... ... .lvficmms Limbattts ..................•. , . .. .. ... MicnmlS nebularis .. . ..... ........... •. ....... Mic r11r11S nigrocinctus ....... ..... •..•. ......... MicrtmlS pachecogili ........ ................... Micr111w proximans ................... , ..... ... 1vficmrus tmer .......................... , .. ... Genus Hydrophis .. ............... ..... ......... Hydrophis platurus ...... , .... ... ......... .... . Family Natricidae............ ........... .... ... ... Genus Adewphis .... •.... .... . .... ...........•.. Adelophis copei ............ , ..... , ............ Adelophis foxi ..........•........... , ......... Genus Nerodia ...........• , .... , .. ........ .... . Nerodit1 e,]'throgmter ... , .•.... ............ .... . Nerodia rhombifer ... ...... • .. ......... ........
293 . 2 96 297 . 2 97 298 299 299 299 300 300 301 3or 302 302. 303 303 304 304 304 305 305 305 308 308 309 309 312 312. 312 313 313 316 317 317 319 319 320 320 321 322 324 324 325 328 328 329 332 332 333 333 336 337 337 338 338 339 340 340 343 343 345 345 345 345 346 348 348
8
Contents Genus Storeria ..... .......... ........... ....... 349 Storeria dekayi .... .... ..... ............... .... 349 Storcria hidalgoemis ..... ........... ........ .... 352 Storeria srorerioides ...•.. .... .............. .... 353 Genus Thamnophis ............ .............. .... 353 Thamnophis bogerti ......... ................... 357 Thamnophis chrysocephalw ..................... , 357 Thamnophis conanti ......... ........ .•...• .... 360 Thamnophis C)'rtopsis ..........•... • . •.•........ 360 Thamnophis elegans ........................... , 361 Thamnophis eques .......... . ......... • ... . . . .. 361 Thamnophis erram ...•.................... .. .. 365 Thamnophis exs11l .. • ...• ...... . . .... ........ .. 365 Thamnophisfidims ....... , . ....•..... . ... •.... 365 Thamnophis godmani .....•..... ........ ....... 368 Thanmophis hammondii .......•. , ..... .. . ...... 368 Thamnophis linen· .... ....• ...• ........ ........ 369 Thamnophis marcianus ............ ... ........ .. 369 Thamnophis melanogasrer .. . ...•........ .•...... 372 Thamnophis mendax ........•...•...... ... •.... 373 Thamnophis nigronuchalis .... ...... ..... ........ 373 Thamnophis postremus ... .... ........... ....... . 376 Thamnophis proximus ... .... ................... 376 Thamnophis pulchrilatus . ....................... 377 Thamnophis rosSllumi . • . .......... . .. . ... . ..... 380 Thanmophis scalaris ........ .............. ...... 380 Thamnophis scaliger ........ .................... 381 Thamnophis sirralis ........ ......... ..... ...... 381 Thamnophis swnichrmti ....... ............ ... . . 384 Thttmnophis 1milabialis .. ... .................... 384 Thamnophis validtts. ....... .............. ...... 385 Family Sibynophiidae .......... .. ............ ...... 388 Genus Scaphiodontophis ...... ..•.. ..... • ...•.. ... 388 Scaphiod,mtophis 1111mtla111s . . .............. ...... 388 Family Viperidae ....... ............ ........ ...... 392 Genus Agkisrrodon ......... . ............•. ... ... 392 Agkistrodon bili11eat11s .. ..•. ..... ...... .•. ...... 393 Agkistrodon co11tortrix ..... .........•...•. ..•... 394 Agkistrodon russeolus . ..... .......•.....•. ...... 395 Agkistrodon taylori ........ ............... ...... 395 Genus Atropoides .................•..•.......... 396 Atropoides mexic111ms ... ... .•...• .•...•.. ....... 397 A11·opoides nummifer .......•...•......... , ..... 397 Atropoides occiduus ............................ 400 Aeropoides olmec ....•......................... 400 Genus Bothriechis .... ...... ... .................. 401 Bothriechis aurifer .......................... , .. 401 Bothriechis bicolor ....................... , . .... 404 Boehriechis rowleyi .. ...... ........... ... ...... . 404 Bothriechis schlegelii ............................ 405 Genus Bothrops ........................ , ........ 405 Bothrops asper ..... , .....•..... , . ............. 408 Genus Cerrophidion ............................. 408 Cerrophidion godmani ...........•. ...... ....... 408 Cerrophidion petlalcalemis .... :...•............. , 412
C e1Tophidion emtzilormn . . .... .. ..•............. Genus Mixcoatlur ........ , ..... ...•............. Mixcoatlur br1rbouri .......... , ..• , . , ..•.... ... /Vfixcoatlus broumi ...............•.• , ..•....... Mixcoatlus mclamm1s ..... ...... .• .....•....... Genus Ophryncus .................•.•......... , . Ophrya cus 11nd11!.ntus .................•.•. ...... Genus Porthidium ... .................•.•.... ... Porthidium dunni ....•........•.....• , •....... Porthidiwn hespere .......................... , . Potthidittm na.sutum ................ , ..•..... , . Porthidium )'UCatanicum ......... .......•.... ... Genus Crotalus ..................... , . , . , .... ... Crotalus aquilw ....................•.... , .. , . Crotalur armst1-o,1gi ....................... , •... Crotalus atrox ........... ...... ......... ...... Crotalus basiliscus .........................•... Crotalus campbelli ........................ , •... Crota!us catalinensis ...................... , .. , . Crotalus cermtes ............. , ............ , .... Crotalm cttlminatus . , .. , .. ............... ...... Crota/11s enyo ............... , ... . ............. Crotalus ericsmithi ...........•........ , .. ..•... Crotalw esteba.nensis .................. , .. , .....
Crotalw intermedius Crotalw larmomi ......... , .................... CrornLus Lepidus ..........•..•.. , .............. Crotnlus mitchellii ....................... , .... . CrotaLus molosrus .........•.. , .. , ........ , ..... Crotalus momlus .........•.. , ..•.............. Crotalus oreg111111s ............................. . Crotr1lus omatus .........•.................... Cmtalus polystictus ......................... , .. Crotalus pricei .......... , .•.... , ...•.......... Crotalus pusillus ...........................•.. Cro ta !us ravus .•. ... ....... • .• .... . ......... .. Crotalus mber .... , ........................... Crotalus scutularus ............................. Crotnlus simus .............•.................. Crotalus stej11egeri ..........•.................. Crotalus tancitarensis ........................... Crotalus tigris ...... , ...... , ..... , ........... , Crotalus tlaloci ..................... , ....•.... Crotalus toto11aws ...... ......... ......... •.... Crotalus transversus ........ .· .......... .... •.... Crotalw triseriatus ............................ , CrotaLus tzabcmz ....•..................... , ... Crotalus viridis . ..........•.• ................. Crotalus willardi .... . .•........ .....•....... .. Genus Sistrurus ................................ Sistmms tergeminus ..........• , .... , , .... , , .. ..
Disuiburion Maps ...................... , .. , ..•.. , .. .. Glossary .............................. , ....... ...... Lireracure Ciced .. , .................... ....... • ....... Index , .. , .. , .....•.. , .......................•. ••••
412 413 413 416 4r6 418 418 4 19 4'9 ,µ2 422 423 426 430 431 431 434 435 435 438 439 442 442 443 443 446 447 450 451 454 455 455 458 459 462 462 463 467 470 470 471 471 474 475 475 478 479 479 487 495 495 504 518 522 557
9
Foreword by Larry David WILSON
The year in which [ am writing chis foreword (2-015) marks the 70"' anniver sary of rhe appearance of one of rhe most seminal pieces of work on Me xican herpetology, che 1945 check.list and key co che snakes of Mexico by Hobart M. SM1m and Edward H. TAYLOR, which appeared in rhe Bulletin of rhe Unired Srares National Museum (now the Narional Museum ofNatural Hisrory). This work of239 pages documented the scare of knowledge about rhis group of squamares exisring at the rime of rhe end of World War 11. This publication was part of a series of duee, the other rwo ccearing of rhc amphibians of Mexico (1948) and the reptiles exdusive of the snakes of Me xico (1950). Thar the 194> checklist and key is still of importance in herpe tology today is anested co by the ciration of this reference in the recenrly publishcd WALLACH et al. compendium Snak1's ofthe World: a catalogue of living and extinct species (201-1). For a number ofdecades, I had a well-worn copy ofthe original until I gave it to a former student ofmine. Now, l have an elecrronic copy l can use as reference for writing rhis foreword. le is not quire che same as having rhe paper original. The liceramre on Mexican snakes since 1945 is voluminous. From a raxo nomic viewpoint and based on che posirions caken by the author of rhis book, the creators ofrhis literature have been responsible for adding a total of46 species and Tl genera to d1e numbers recogni7.ed by SMITH and TAVl.OR (1945). As expected, these figures include all che synonymizations and resur rections, as well as the description of newly-discovered taxa at boch the spe cific and generic levels. Even though I have chosen co compare Peter HElMES' Mexican snake book co the checklist and key ofSM1TH and TAYLOR (c945), chece are many distinc rions in chc two works chac make Heime.s' work fur more useful than its pre decessor, as is to be expected, given the span of time between the publica tion of chcsc rwo works and the advances that herpetological publishing has undtrgone. The most obvious distinccion is that rhe H1:1MES' book is illust rared wirh a large number ofexcellent photographs ofche snake species, many of chem cakcn by rhc amhor himself. Jncersperscd among chem are a ntLm bcr of habicac phorographs illusrraring the kinds of places in which give11 species can be found. Even so, the author acknowledges the assistance of nu merous individuals who invited him onro their fieldcrips so chat he had a chance to gacher the photos of his used in chis book. Thus, even rhough, in rbis age of multi-authored sciencific works, HEJMES is che single author of rhis book, ic is comforting to see chat he was helped along by a subscancial number of people. The photographs are nearly all in color with the snake placed ag.unsc a natural or naturalistic background, In some instances, a given species is illustrated by more than one photograph: rhis is especially the case when che juveniles have a different parcern chan do the adults (c. g., Ph1p101111x poeci/011otm) or the species illusrraces more rhan one pacrern rype (e.g., Sonom michoacanensis). For users who are reasonably familiar with snakes, du:" phorographs alone might suffice co allow for idcnrific.1cion of rhe species in question, which identification can then be checked againsr in formation in the appropriate key and the wrinen accoum of the species. ft appears likely rhac che phorographs alone are worth the price of the book. Of great use also in the idenrification of che snakes included are the dis tribution maps included in a single secrion after the species accounrs. l think rhe reader would find ic useful ro read che seccion on rhe natural environment ofMexico chat precedes the keys and species accounrs in order co become fa miliar wirh the physiographic makeup of chis counrry, which makeup shapes rhe distributional patterns evident among these snakes. Thus, terminology for areas such as chc Sierra Madre Occidencal, Sierra Madre Oricncal, lsrh mus ofTehuantepec, and Yucatan Peninsula will have a chance ro become familiar enough ro the reader such chac che descriptions of geographic ranges of che Mexican snakes in che species accoums can help envision where they occur, which undemanding can chen be checked against rhe distribution maps included. Dr. HEIM ES includes a briefdiscussion ofche major distributional panerns of Mc.xican snakes based on the face char elemenrs ofborh the Nearctic and Neocropical biogeographical realms are presenr in d1e counrry. The cwo realms are separated, of course, by rhe Tropic of Cancer, which crosses the counrry from west to cast just north of Mazatlan in souchern Sonora, thence south ern Durango, northern Zacacecas, northern San Luis Potos!, extreme soud1cm
Nuevo Lecin, and, finally, just souch ofCiudad Victoria in southern Tamau lipas. The aurhor divides che snake fauna into four groups, depending on how cl,e componem genera are distributed relative co chis line of division. The four groups include che 11 genera chat me purely Nearcric in dimibu rion, chose 44 that are completely or almost completely Ii.mired ro che Nco rropicaJ realm, chose 3r che disrriburion ofwhich occur subscamially in both regions, and, finally, che cwo genera rhac are found in tropic.ti and subtropi cal regions globalJy, one ofwhich occurs natively along the Pacific coast (Hyd rophis platurus) and rhe other of which is introduced widely in che country from the Old World (/T1dot;rphlops bmminw). The first ofthe dichotomous keys is co che 90 genera of Mexican snakes, which occupies rwo and one-halfpages. This key is rhe one rhar will be put to greatest use by che reader. Such keys consisr of a series.ofcouplers, which, when employed, allow che user to proceed in a seep-wise fashion hopefully co a generic name rhar will belong co the animal in hand. Using such keys requires a bit of practice on rbe pare of the user, but a great deal ofskill on the pare ofrhe composer. Once a given genus is reached, then the reader is directed co che portion of the guide where char genus is discussed. Ofren, another key is encountered in those instances in which the genus in quesrion contains more rhan one species resident in Mexico. The bulk of chis book consists of the accounts of che various caxa in volved in the systematic arrangement ofthe Mexican snakes, beginning with rhe infraorders and cominuing on through rhe families, genera, and species. For each of che caxa above the species level, rhe author presems a brief sum mary of their discciburion, appearance, and habits. As expected, rhe major iry ofthis pare ofche book is comprised ofrhe species accounts, one for each of the 390 species included; The species accounts follow a standard format, including chc snake's scientific name and aurhorship of chat name, a lisc of common names in English, Spanish, and occasionally an indigenous lan guage. This secrion is follow�d by a briefsynonymy for che species, a shore identification section, a more-detailed descripcion, a detailed sratement of discribucion (including a listing ofsources for distributional information), and a scacemenr of natural history informadon (also including references documenting chis information). The amount ofinformation the amhor bas packed in each of these accounts consricurcs a stunning achievement char would seriously ta.-x the energies ofa team of researchers, ler alone a single in dividual. This attention to decail is what will give chis work value long into rhe present century for both snake aficionados and professional herpetolo gists alike. As noted above, che distribucion maps are placed after che species accoums and precede a glossary oftechnical terms that spans about a page and a half. The final portion of rhe texc of rhe book is composed of an amazing com pilation ofliterarnre percaining ro Mexican snakes char occupies 34 pages and includes references published throughour che span ofca.xonomic work, i. e., since che publicarion ofCarolus LINNAEUS' masterpiece in 1758 all rhe way co rhe current year. This compilation will prove co be a goldmine ofinfor macio1i of significant use ro specialises, especially younger ones who mighr nor have gained yet the experience chat older workers have had ro spend decades co obtain. 1 am sure that chose who realize whar has to go inco a piece of work like this book are going to be hugely impressed with Dr. HEIMES' accomplish ment. First, he spent seven years (1997-2004) in Mexico gathering the data chat form the basis of che book. He also spent several years assembling this 570-page volume..Finally. Peter spent several months ofback-and-forch with the publisher as he worked ro keep his book current with che sizable m11n ber of changes in snake sysremaci� in general and Mexican ophiology in particular char have occurred apace. The amount of dedication required to pull off chis achievemenr is highly admirable. [n preparing chis foreword, it has become abundanrly evident to me char Peccr Hw,rns' book ro rhe snakes of Mexico will establish a new srandard of excellence for rhis kind ofwork chat will be the envy of all rhose wbo are enrhralled by che study ofchis marvelous group ofcreatures in tbc endlessly fascinating country of Mexico. For those who mighr seek co equal or becrer Peter's accomplishment, chey arc going co have to work exrn:mely hard co be able to clear the bar established by him in chis work.
10
Introduction
The herperofauna of Mexico is one of the richest in the world. hoRES-VILLELA and CANSECO-MARQUEZ (2004) recognized u65 species of amphibians and reptiles for the country, and chis num ber is growing rapidly with the discovery of new species. For example, FLORES-VILLELA (1993) listed 291 amphibian and 706 reptile species, but only cen years later these numbers had in creased to 36r ( + 24.0%) and 804 ( + 13.8%), respectively (hoRES ViLLELA & CANSECO-MARQUEZ 2004). Major factors contributing co Mexico's biological diversity are the exceedingly rugged and divergent topography of th.is country and its location on the bor der between temperate and tropical zones. Mexico is also located near the border between two continents that are connected by a narrow land bridge which has allowed a mixing of fauna! elements during earlier and more recent geological periods. This book is a compilation of the widely scarcered information on MeKican snakes, including recent contributions involving their systematics, geographic disrribution and natural history. Over the past cencury and the first decade of the present, a tremendous amoLLnt of literacure has been published on chis subject, including thorough monographs and other eKtensive papers dealing with selected groups of snakes occurring in Mexico (e. g., MAU BER 1940d on Leptotyphlops (now Rena); BLANCHARD r921 on Lampro peltis; ORTENBURGER 1928 on Masticophis; STULL 1940 on Pitu ophis; KLAUSER 1935a, 1940a 011 Phyllorhynchus; Kl.AUBER 1941 on Rhinocheilus; SMITH 1941g on Drymarchon; SMITH 1942f on Adel phicos; TRAPIDO 1944 on Storeria; l:-.;,,.
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'1 ventrals in males and 225241 in females, and 69-86 divided subcaudals in males and 5984 in females. Distribution: 'Ji·imorphodon vilkimonii is an inhabitant of the Chihuahuan Desert, ranging from the Rio Grande drainage sys tem of New Mexico and the Big Bend region ofTexas southward inro northern Chihuahua. The vertical distribution extends from about 900 co 1,850 m (SCOTT & McDIARMID 19846, WERLER & DrxoN 2000, LADuc &Jot:1.NSON 2003, LEM0s-Esr1NAL &SMITH 2007a, DEVITT et al. 2008, GATICA-COLJMA et al. 2012). Natural history: This species is most commonly found in dry and rocky mountainous areas of oak-pine woodland, but also occurs in intervening floodplains and shallow canyons (WERLER & DrxoN 2.000, LEMOS-EsPINAL & SMITH 2007a). k is active pri marily ar night, seeking refuge during the day under rocks, in cre vices among rocks, or sometimes in caves. Ir preys chiefly on lizards, which preswnably are detected asleep in their rock-crevice shelter; small rodents, bars, birds and snakes are also eaten. Ma ring presumably occurs in autumn. TENNANT (1984) reported that a captive female layed 6 eggs in late June. The incubation lasted 77 days and the hatchlings measured an average of 216 mm TL.
Species Accouncs: AJerhinophidia: Dipsadidac
Family Dipsadidae
BONAPARTE,
1838
This is one of the largest snake families, wich all representatives restricted to the New World (VmAL et al 2010, G R.AZZIOTIN et al. 2012). According to PYRON et al. (2013), che Dipsadidae com prise 95 genera and 738 species, of which 28 genera and 127 species are fow,d in Mexico. Dipsadids ;i.re primarily tropical, with most occurring in Central America, South America and the West Inrues. They vary gre;i.tly in body size (up to 280 cm TL) and in ecology. The family includes rather unspecialized terrest rial species that live within the forest floor litter, small burrowing snakes with riny eyes and a pointed snout chat feed mostly on earthworms, others that dig up toads with their spade-like snout, colorful coralsnake mimics char prey on ocher snakes, and slender semi-arboreal species char are specialized predators of sleeping anole lizards, frogs eggs, or slugs and snails. Tretanorhinus nigro Luteus and some other inhabitants of lakes, swamps and sluggish sffeams are rarely seen our of the water. The family Dipsarudae is similar to the family Colubridae in mosc morphological fea tures, although the former contains a higher percentage of rear fanged species, some of which possess fairly potent venom. So far as known all dipsadids lay eggs.
Genus Adelphicos JAN, 1862 This genus contains eight species of small fossorial snakes that collectively range on the Atlantic versanr from sourhern Tamau lipas to northern Honduras, and on the Pacific versant from west ern Oaxaca to Guatemala (CAMPBELL & FORD 1982, CAMPBELL & BRODIE 1988, SMITH et al. 2001); five species are found in Mexico. These highly secretive snakes occur in forested habitats from sea level up to approximately 2,900 m. Members of the genus are characterized by the following suite of fearures: body moderately slender to moderately stout, cylind rical; rail shore; head small, conical, nor or only little wider than neck; eyes small and with a round pupil; dentition aglyphous, with maxillary teeth similar in size; basic colubrid complement of enlarged head plates; dorsal scales smooth and in 15 rows through out length of body; subcaudals divided; anal plate divided. Key to the Mexican Species of the Genus Adelphicos ra. Third infralabial weU-developed and subequal in size to second; chinshields nor grearly expanded toward lip ...................... 2 16. Third infralabial absent, or greatly reduced in size and con fined to labial border; chinshields greatly expanded toward
lip .................................................................................... 3
2a. Subcaudals more than 36; rniddorsal srripe involving para verrebral scale row; subcaudal surface generally suffused with black ................................................ Adeiphicos !atifasciatus 26. Subcaudals fewer than 37; rniddorsal stripe, if present, in volving vertebral row only; a black median subcaudal stripe ............................................................ Adelphicos nigriLatttS
?.Of
3a. Chinshields separated from lip by a very narrow third labial; subcaudals 36 to 49; belly frequently heavily pigmented ...... . ................................ .... ....... ................. Adelphicos visoninus 36. Chinshields bordering lip, third infralabial absenc; belly not or lirtle pigmented ............................................................ 4 4a. Subcaudals 24-35; ground color very dai:k, lines scarcely vi sible; never any pigment on belly, subcaudal surface profusely pigmenced ................................................. Adelphi cos sargii 46. Subcaudals 31-50; lines on sides and back distinct; belly some times with some pigment; a black median subcaudal stripe ..................................................... Adelphicos quadi-ivirgatus Adelphicos Latifasciatus LYNCH & SMITH, 1966
Oaxacan earth snake; ocotera de Oaxaca
Adelphicos veraepacis latifosciatus LYNCH & SMlTH 1966: 66. Type locality: "Sierra Madre north of Zanatepec, Oaxaca, Mexico".
Adelphicos latifosciatus CAMPBELL & FoRD 1982: 13.
Identification: A small, fossorial snake wich a dorsal pattern of three dark stripes; middorsal stripe i_11volviJ1g paravercebral scale row; third infralabial well-developed; chinshields noc expanded coward lip; dorsals in 15 rows; subcaudals more chan 45 in males and more than 36 in females. Description: TL up to 44 cm. The body is moderately robust, the cail is moderate in males and shore in females. The dorsal coloration is pale to reddish brown with three dark brown stripes. The middorsal stripe covers the vertebral and adjacent halves of tl1e paraverrebral scale rows, the lateral stripe involves scale row 3 and adjacent pans of rows 2 and 4. The head is dark brown or reddish brown above including the upper parts of the supralabi als. The gular region and ventrals are cream-colored and immacu late except for pale mottling on the outer portion of the vencrals. The subcaudals are pale with a dark midventral scripe along their median sutures. There are 7 supralabials, 7 infralabials, a single elongate scale in the loreal-preocular area, 2 postoculars, 1+1 tem porals, 125-128 ventrals in males and 133-138 in females, and 465r divided subcaudals in males and 37-41 in females. Distribution: Adelphicos latifosciatus is known from the north western pare of the Sierra Madre de Chiapas in extreme western Chiapas and adjacent southeastern Oaxaca. The vertical distri bution is from about 1,500 to 2,000 m (CAMPBELL & FoRD 1982, CAi\1PBELL & BRODIE 1988, JOHNSON 1989). Natural history: AdeLphicos fatifasciatus occurs in cloud forest and pine-oak forest. le is more active below ground than on the surface and is usually found in deep leaf litter or under rocren logs. Like ocher members of the genus, ir presumably feeds on soft-bodied invertebrates and lays eggs.
202
Species Accounts: Alerhinophidia: Oipsadidae
Fig. 245: Adelphicos latifasciatus, La Sepulrura, Municipio de Cimalapa, Sierra Madre de Chiapas, elevarion 1,200 m. A. RAMIREZ VELt\ZQUEZ
Fig. 246: Adelphicos nigrilatus, Mirzir6n, Chiapas. C. GRUNWALD &
J. }ONES
Fig. 247: Adelphicos nigrilatus, San Crisr6bal de las Casas, Meseca Cenrral de Chiapas, elevation 2,300 m. P. HEIMES
_ Species Accounts: Alerhinophidia: Dipsadidae
203
Fig. 248:
Adelphicos quadrivirgatus, Caternaco, Sierra de Los Tuxdas, Veracruz.
P.
HEIMES
Fig. 249:
Adelphicos q11adrivirgat11s, Zacapoaxrla, Puebla.
P.
HEIMES
Fig. 250:
Adelphicos visoninus, ANP El Zapocal, Tuxda Gutierrez, Chiapas, elevation 650 rn. A. RAMIREZ VELAZQUEZ
Species Accounts: Alerhinophidia: Dipsadidae
204
Adelphicos nigrilatus
SMITH, 1942
Chiapan earth snake; ocotera de Chiapas
Adelphicos veraepttcis nigrilatus SMITH 1942f: 182. Type locality: San Crist6bal de las Casas, Chiapas, Mexico. Adelphicos nigrilatus CA-"APBELL & FoRD 1982: ro. Identification: A small fossorial snake with a dorsal pattern of dark stripes; middorsal stripe, if present, involving vertebral scale row only; third infralabial well-developed; chinshields not ex panded coward lip; dorsals in 15 rows; subcaudals fewer than 37 in males and fewer than 27 in females. Description: This relatively scout species reaches a maximum TL of about 30 cm in males and 45 cm in females. The dorsal ground color is bright orange, dark red or pale brown wirh a par tern of 2, 4 or 5 dark stripes. The middorsal srripe involves rhe vertebral scale row only; in many individuals it is absent or bro ken. The vertebral scale row is pale in coloration when the srripe is absent. The lateral stripe usually is rhe most distinctive one and involves 1-3 scale rows. The paraverrebral stripe, if present, is less conspicuous than the lateral stripe and usually involves one scale row. The veneer is much paler rhan rhe dorsum, only a few specimens possess a heavy pigmentation midventrally. A black midvencral stripe is present on the posterior portion of che body and on the rail. There are 7 supralabials, 7 infralabials, a single elongate scale in the loreal-preocular area, 2 posroculars, 1+1 temporals, 113-128 ventrals in males and 123-135 in females, and 26-36 divided subcaudals in males and 21-26 in females. Distribution: Mesera Cenrral of Chiapas. Adelphicos nigriltttus is known from rhe vicini ty of San Cristobal de las Casas south ro the Guatemalan border at 2,200-2,900 m elevation (CAMPBELL & FoRn 1982, CAMPBELL & BRODLE 1988, ]oHNSON 1989).
Natural history: This secretive snake lives underground, in piles of plant rubbish or under rotten logs in pine-oak forest and pine-oak-Liquidambar forest QoHNSON 1989). le feeds on earth worms and other soft-bodied invertebrates (CAMPBELL & FORD 1982). Adelphicos quadrivirgatus ]AN,
1962
Description: TL up co 40 cm. The dorsal ground color is grayish brown, medium brown or red. There usually are four black nar row black srrjpes, but these may be obfuscated by a dark ground color, especially in the subspecies A. q. newmanorum. A dark dorsolareral stripe is present on scale row 2 and portions of the adjacent rows, and a dark vencrolateral stripe occupies scale row 5 and portions of the adjacent rows. The central portion of the vemrolareral stripe is often pale, creacing two rhi.n Lines. The mid dorsal area is usually slightly darker than rhe ground color. The rop of the head is dark brown. The veneer may be cream, pale yellow, pink or salmon. In some individuals rhe ventrals are im maculate, while in ochers they are heavily peppered wich dark pigment. A dark midvenrral line on the rail often extends for ward co rhe last vencrals. T here are 7 supralabials, 7 infralabials, a single elongate scale in che loreal-preocular area, 2 posroculars and 1+1 temporals; the anterior chinshields are greatly expanded and reach the lips; there is no chird infralabial and the second is greatly reduced; chere are 124-155 vencrals (124-142 in A. q. quadrivirg;attts and 141-155 in A. q. newmanorum) and 31-50 divided subcaudals. Distribution: Adelphicos quadrivirgatus occurs at low and mode rate elevations (up to 1,500 m) on rhe Atlantic versanr from southern Tamaulipas and adjacent areas of Nuevo Le6n to the Sierra de Los Tuxclas of southern Veracruz. Two subspecies are recognized: A. q. newmanorum occurs in the northern pare of rhe range from Tamaulipas sou ch co San Luis Potosi, and A. q. quadrivirg;atus is found from Hidalgo southward (MARTIN 1955, 1958, SMLTli 1942f, CONTRERAS-A.RQUIETA 1989, PEREZ-HIGA.REDA & SMITH 1991, CANSECO-MARQUEZ et aL 2000, SMITH et al. 2001, MENDOZA-Qu1JANO et at. 2003, 2005, LAZCANO et al. 2004, 2005, D1xoN & LEMos-Esr>INAL 2010, Le.Mos-ESPINAL & DocoN 2013, FARR et al. 2013). Natural history: This terrestrial and fossorial snake inhabits tro
pical deciduous forest, lowland and monrane rai_n forest, cloud forest and pine-oak forest. Tc is nocturnal and spends most of its rime underground, under rorren logs or in deep leaf litter at the base of large trees. Jt feeds predom inanrly on earthworms. One to five eggs are laid during the early rainy season in May or June (MARTIN 1955, CAMPBELL 1998a). In southern Veracruz, adurch of three eggs was found in a termite nest some 1-1.5 m above ground; one hacchling measured 86 mm in TL (PEREz-HrGAREDA & SMITH 1989).
Mexican earth snake; zacatera
Rhttbdosoma lineatum GUNTHER 1858: rr. Adelphicos quadrivirgatum ]AN r862a: 18. Type locality: "Mexico", restricted to Jicaltepec, Veracruz, Mexico, by SMITH & TAYLOR (19506). Adelphicos quadrivirgatus quadn'virgatus SMITH 1942f: 188. Adelphicos quadrivirgatus newmanorum TAYLOR 1950: 442. Type locality: Xjlirla, San Luis Potosi, Mexico. Identification: A small fossorial snake with a dorsal pattern of 4 dark stripes; central portion of venrrolateral stripe often pale; third infralabial absent; chinshields greatly expanded and reach ing Jip; dorsals in 15 rows.
Adelphicos sarg ii (F1SCHER, 1885) Sargi's earth snake; zacarera de Sargi
Rhegnops sarg;ii F1SCHER 1885: 92. Type locality: Guatemala. Adelphicos sargii CoPE 1887= 85. Adelphicos quadrivirg;atus sargii SMITH 1942f: 192. Identification: A small fossorial snake with a pattern of 5 or 7 obscured stripes on a dark brown ground color; third infralabial absent; chinshields greatly expanded and reacl1ing lip; dorsals in 15 rows; subcaudals fewer chan 36.
Species Accouncs: Alechinophidia: Dipsadidae
Description: TL up to 31 cm. The dorswn is dark brown with 5, sometimes 71 greatly obscured stripes, including a verrebral stripe char is barely visible in some individuals; a narrow dorsolareral scripe extending along rhe adjacent edges of scale rows 4 and 5; a.nd a broad ventrolateral stripe. The central portion of the ventro lareral stripe is sometimes pale, creating two thin lines. The ventral surfuce of head and body are pale and immaculate, the subcaudals are profusely stippled with dark pigment. There are 7 supralabials, 5-6 infralabials, a single elongate scale in the loreal-preocular area, 2 posroculars and 1+1 temporals; rhe anterior chinshields are greatly expanded and reach the lip; there is no tl1ird infralabial and rhe second is greatly reduced; there are 125-147 ventrals and 2435 divided subcaudals. Distribution: On rhe Pacific versanr from rhe Soconusco region of extreme southern Chiapas ro southwestern Guatemala (S M ITH r942f ). Natural history: This secretive snake occurs in tropical lowland and monrane rain forest. Adelphicos visoninus (COPE, 1866) Middle American earth snake; zacarera de cuarro lineas Rhegnops visoninus CoPE 1866a: 128. Type locality: Honduras. Adelphicos visoninus C OPE 1887: 85. Adelphicos quadrivirgatus visoninus SM rTH 1942f: 186. Identification: A small fossorial snake with a dorsal pattern of 4 or 6 dark stripes; infralabials 2 and 3 very narrow; chinshields greatly expanded and almost reaching lip; dorsals in 15 rows; sub caudals more than 35.
205
Natural history: This secretive, burrowing snake occurs in tro pical deciduous forest, lowland and montane rain forest, cloud forest and pine-oak forest. Ir is predominancly nocturnal and feeds on earthworms and ocher soft-bodied i.nvenebrares (LA.Noy et al. 1966, SEIB 1985, CAMPBELL 1998a). Eggs are laid during the early rainy season.
Genus Amastridium COPE, 1861 This is a genus of rwo moderate-sized species, one occurring in Mexico and upper Central America, and the other in lower Central America. These are relatively uncommon terrestrial and semi fossorial snakes that inhabit the leaf litter of undisturbed humid forests. Perhaps rhe most obvious feature of rhe genus is a dis tinct canrhal ridge rhat makes their head shape look like that of a viper. Ocher defining characteristics include a moderately slen der, cylindrical body with a moderately long rail; head noticeably distinct from neck; eyes large and with a round pupil; dentition opistoglyphous, with enlarged rear maxillary reerh; basic colubrid complement of enlarged head plates; dorsal scales smooth and in 17 rows throughout length of body; supracloacal tubercles in males; anal place and subcaudals divided. Amastridium sapperi (WERNER, 1903) Sapper's rusty-headed snake; culebra cabeza de cobre de Sapper Mimometopon sapperi WERNER 1903a: 349. Type locality: Guatemala. Amastridium sapperi DUNN 1924: 1. Amastridiurn velifenan W1LSON & MEYER 1969: 145. Amamidium velifemm sapperi SMITH r97ra: 255.
Description: This relatively slender species reaches a maximum TL of about 40 cm. The dorsal ground color is pale gray, pale ro dark brown, or reddish brown. There are usually four narrow black srripes, bur these may be obfuscated by a dark ground color. A middorsal srripe is absent or very poorly defined. A dark brown or black dorsolareral stripe is present on scale row 5, or on rhe adjacent halves of scale rows 4 and 5. A dark ventrolateral stripe is present on scale rows 1 and 2, sometimes including the lower half of row 3. The central portion of the ventrolateral stripe is often pale, creating two thin lines. The top of rhe head is dark brown. The belly is cream, pale yellow, pink or salmon. In some individuals the ventral surfaces are immaculate, in others they are heavily stippled with black pigment. There are 7 supralabials, 7 infralabials, a single elongate scale in the loreal-preocular area, 2 postoculacs and 1+1 temporals; the anterior chinshields are greatly expanded and rhe second and third infralabials are very narrow, barely excluding rhe chinshields from the labial border; there are 126-148 ventrals and 36-49 divided subcaudals.
Identification: A small to moderate-sized snake having a uni formly dark dorsum and a lateral series of pale small dots; top of head usually reddish brown with dark streaks; dorsals in 17 rows.
Distribution: On rhe Aclanric versanc from che Sierra Juarez in nonheascern Oaxaca southward through extreme southern Vera crnz, Tabasco, and northern and eastern Chiapas ro northern Guatemala and Honduras. The vertical distribution ranges from near sea level to about r,600 m (S MITH r942f, ALVAR£Z DEL TORO 1973, 1982, SMITH et al. 2001).
Distribution: On the Atlantic versanr from Nuevo Leon and Tamaulipas southward co western Honduras, exclusive of rhe Yucatan Peninsula, and on the Pacific versant from extreme south eastern Oaxaca to southwestern Guatemala. Amastridium sapperi appears to have a discontinuous distribution throughout irs range. In Mexico, it is known from scattered localities in west-cent-
Description: TL up to 73 cm. The dorsal coloration is rather uniform dark olive, dark brown or nearly black, with a series of wh.jtish or pale yellow docs on scale rows 5 or 6; the docs are sep arated from each other by 1-5 scales. The cop of the head is red dish brown or copper with irregular black streaks; the conspicuous reddish head color may fade co a dark yellowish brown in large specimens. The dark dorsal color extends from the body forward to the neck where it runs our in form of a finger or wedge that reaches about ro the parietals. A dark brown postocular stripe ex tends from che eye co the angle of che jaw or slightly beyond. The venter is paler than the dorsum and heavily mottled with gray or gray-brown pigment. There are 7 supralabials, 9 infralabials, 1 loceal, I preocular, 2 poscoculars, 1+2 temporals, 7 supralabials, 9 infralabials, 144-170 ventrals and 77-86 divided subcaudals.
206
Species Accounts: AJerhinophidia: Dipsadidae
Fig.
251:
Ammtridittm sapperi,
Valle de Jalpan, Queretaro. R.W. HANSEN
Fig.
252:
Chersodromus liebmanni,
Cordoba, Veracruz. P. HEIMES
Fig. 253:
Chersodromus mbriventris,
Chilijapa, Tepehuacan de Guerrero, Hidalgo.
C.
BERRIOZABAL-ISLAS
Species Accounts: Alethinophidia: Dipsadidae
Fig. 254: Clelia scytalina, La Fortaleza, Municipio de Santa Maria Chimalapa, Oaxaca.
Fig. 255: Clelia scytalina, juvenile, Ejido Lazaro Cardenas, Las Choapas, Veracruz.
207
T. DEVITT
L. CANSECO-MARQUEZ
208
Species Accoums: Alerhinophidia: Dipsadidae
ral Nuevo Leon, somhwesrem Tamaulipas, northern Quereraro, northern Hidalgo, norrhern and southeastern Puebla, southern Veracruz., southeastern Oaxaca, and western and northeastern Chiapas. The vercical distribution extends from near sea level co around 1,520 m. Distribution informarion is given in DuNN 1924, SMITH 1944, M ARTIN 1955, SMITH 1971a, BLANEY & BLANEY 1978a, WILSON 1988a, PEREZ-HIGAREDA & SMITH 1991, LEE 1996 and CANSECO-MARQUEZ & GUTIERREZ-MAYEN 2006b. Natural history: This relatively uncommon diurnal and crepus cular snake inhabits the leaf liner zone of evergreen and semi evergreen seasonal forest, tropical rain forest and cloud forest. Irs diet consists of small lizards, frogs and myriapods (M ARTIN 1955, CAMPBELL 1998a). Ir is oviparous.
Genus Chersodromus
REINHARDT, 1860
This genus is composed of rwo poorly known species of small blackish snakes that are terrestrial and semifossorial. They are dis tribmed in the Sierra Madre Oriental and Sierra Madre de Oaxaca of eastern Mexico. Several authors have suspected char Cherso dromus is most closely related to the genera Ninia and Geophis (DUNN 1935, TAYLOR 1949, DOWNS 1967). Members of the genus .tre cbaraccerized by the following com bination of fearures: body slender and cylindrical, rail medium in length; head small and slighdy distinct from neck; eyes rela tively small and with a round pupil; dentition aglyphous, with maxillary reeth similar in size; basic colubrid complement of en larged head places, with exception of fused prefrontals; dorsal scales smooth or weakly keeled and in 15 or 17 rows at midbody; subcaudals divided; anal place undivided. Key to the Species of the Genus Chersodromus Dorsal scales in 17 rows ....................... Chersodromus liebmanni Dorsal scales in 15 rows ...................... Chersodromus riibriventris Chersod1·omus liebmanni REINHARDT, 1860 Liebmann's earth runner; corredora de Liebmann Chersodromus liebmanni REINHARDT 1860: 242. Type locality: Mexico; resrricred to Cuaurlapan, Veracruz, Mexico, by SMITH & TAYLOR (19506). Chersodromus nigricans REINHARDT 1860: 245. Opisthiodon torquatus PETERS 1861a: 461. Type locality: Huanusco [= Huarusco], Veracruz, Mexico. Dirosema collare WERNER 1900: 197. Identification: A small blackish snake with a pale collar; a single scale in the prefrontal area; mental shield couching chinshields; dorsals in 17 rows. Description: This species reaches a TL of about 30 cm. The dor sal coloration is deep slate-black, extending omo the outer rips of the ventrals. A pale yellow or whitish band with irregular borders crosses the back part of the head involving the posterior half of
the parierals. The pale band passes down onto the gular region where it widens. Head and supralabials are black, with anterior supralabials having pale centers; mentals and anterior infralabials are also black. The belly is cream wirh dark pigmenc increasing coward rhe rail. The tongue is reddish brown with black rips. There is a single large scale in the prefrontal area and a single large scale in rhe loreal-preocular area. There are 7 supralabials, 6-8 infrala bials, I poscocular (ofren very small or fused with the supraocular), 1+2 temporals, 124-.142 vemrals and 32-43 divided subcaudals. Dorsal scales are finely srriared; rhe scales of the first row are vis ibly broader than orher dorsal scales. Distribution: Eastern Mexico. Chersodromus liebmanni ranges in the Sierra Madre Orienral ar moderate elevations (800 co 1,800 m) from northern Puebla and central Veracruz south to northern Oaxaca (PEREZ-HlGAREDA & SMITH 1991, CASAS-ANDREU et al. 1996). Natural history: This terrestrial snake i11habirs montane rain fo resr and cloud foresr. Ir is rarely seen on the surface bur usually is found under rocks aud logs or beneath leaf licrer or piles of de caying vegetation. Nothing is known of its habits; it presumably feeds on invertebrates and lays eggs. Chersodromus mbrivent:ris (TAYLOR, 1949) Red-bellied earrh runner; corredora de pan2a roja Schmidtophis mbriventris TAYLOR 1949: 193. Type locality: "near Xilirla (Xilirla Region), San Luis Porosf, Mexico'', Chersodromus rubriventris D1xoN & KETCHERSID 1969: 163. Identification: A small black snake with a bright red-orange belly and a complete or interrupted pale coUar; a single scale in the pre frontal area; mental separated from chinshields; dorsal scales in 15 rows. Description: TL up co 34 cm. The dorsal coloration is slate-black, extending onto rhe outer tips of the ventrals. A yellowish orange parietal band with irregular borders crosses the back parr of the head and passes down onto rhe gular region where ic widens; it may be broken inro spots dorsally. The head is black anterior and posterior co rbe parietal band; rhe first four supralabials are also black. The throat is cream-colored, the belly and underside of che mil are bright reddish orange, the posterior borders of ventrals and subcaudals are whirish. There is a single large scale in rhe pre frontal area and a single large scale in the loreal-preocular area. There are 6 supralabials, 6-7 infralabials, I posrocular, no anrerior remporal, 125-127 ventrals and 41-43 divided subcaudals. Distribution: Chersodromus mbriventris is known from the Sierra Madre Oriental of southeastern San Luis Porosl and adjacent northeastern Quereraro and northern Hidalgo. Elevation records range from 700 co 1,650 m (TAYLOR r949, P ADILLA GARcfA & PINEDA LOPEZ 1997, DIXON & LEMOS-ESPIN AL 20!0, LEMOS EsPIN AL & DIXON 2013). Natural history: Chersodromus rubriventris ranges from tropical forest through cloud forest up into pine-oak foresr. Two indivi-
Species Accouncs: Alethinophidia: Dipsadidae duals were found under moist coning logs along a karst hillside (DIXON & KETCHERSID 1969, DIXON & LEMOS-ESPINAL 2010). This snake presumably feeds on invertebrates and lays eggs.
Genus Clelia
FITZINGER, 1826
The genus Clelia (mussuranas) conrains eight species of large terrestrial snakes. The distribution of this genus is centered in South America; cwo species are found in Central America, one of whid1 (C. scytalina) ranges northward into Mexico (PETERS & OREJAs-MrRANDA 1970, ZAHER 1996, FRANCO et al. 1997, LEY NAUD & BucHER 1999). The ocher species in Central America is Clelia clelia; C. clelia was reported from several localities in Chia pas and Verdcruz by SMITH (1943a) and subsequently was reported from Mexico by several authors (SMITH & TAYLOR 1945, ALVAREZ DEL TORO 1973, FLORES-VILLELA 1993). STUART (1963) and BAILEY (1970), however, emphasized mac the material described by SMITH (1943a) represents C. scytalina, and SMrTH & PEREZ-HIGAREDA (1989) stared char all references to the genus Clelia in Mexico per tain co chis species. These snakes are powerful conscriccors, bur they also possess venom that aids chem in subduing their prey. The posterior reerh on rhe maxillary are enlarged, grooved, and associated wirh Du vernoy's glands. Mussuranas are predominantly ophiophagous and apparenrly are immune co the venom of venomous snakes which form part of their natural dier. All species of rhis genus undergo a remarkable oncogenecic color change, wirh juveniles having the dorsum red wirh a black head and a black neck sepa rated by a pale collar. Adults are uniform glossy grayish black or bluish black. Ocher features chat characterize the genus include a long and rather thick body with a moderately long tail; a rela tively small head chat is only lirrle disrincr from the neck; small dark eyes with subcircular pupils; basic colub6d arrangemenr of enlarged head places; dorsal scales smooch and in I7 or r9 rows chroughour length of body (17 in C. scytalina and 19 in C. clelia); subcaudals divided; anal plate undivided.
Clelia scytalina (COPE, 1867) Mexican snake eater, brown mussurana; culebrera mexic.'lna, cu lebra negra
Scolecophis scytaLinus COPE 1867: 320. Type localiry : "near Tabasco", Mexico. Oxrhopus proximus BocouRT 1897: 856. Clelia clelia immacu!atus SMITH 1942a: 394. Type localiry: Guada lajara, Jalisco, Mexico (in error, according to ZWEIFEL 1959c). Clelia scytaLina STUART 1963: 90. Identification: Adults have an iridescenr grayish black or bluish black dorsum; juveniles have a red dorsum, black head and pale yellow collar which is bordered behind by black pigmenc; venrer cream-colored and immaculate; dorsal scales in 17 rows. Description: A medium-sized to large snake reaching a maximum TL of about 200 cm; females tend co be larger than males. Juve niles and subadults are uniform red above with a black spot on each scale rip; the middorsal scales have more dark pigment than
209
the lateral scales. The distal half of me upper rail surface is black. A black nape patch is separated from a black head cap by a pale yellow collar that covers parts of me posterior head plates and rhe first 4-5 dorsal scale rows. Adults are slate black or bluish black above with scale rips darker. Juveniles and adults have a cream colored venter. The iris is black. Subadults measuring about 7090 cm in TL usually show a transitional color phase characterized by a black head and brown dorsum. There are 7 supralabials, 79 (usually 8) infralabials, 1 loreal (or none), r preocular, 2- posrocu lars, 2+3 or 2+2 temporals, 202-228 ventrals and 75-92 divided subcaudals. Distribution: Clelia scytalina ranges at low and moderate eleva tions (up to 1,200 m) from Veracruz on the Atlantic versanr and fromJalisco on the Pacific versanc southward through Central America co Colombia (PEREZ-SA NTOS & MORENO 1988, AuTH 1994, SAVAGE 2002). This species is generally rare in Mexico except for rhe Sierra de Los Tuxrlas region in southern Veracruz where ir is considered co be relatively common (PfaEz-HIGARED A & SMITH 1991, VOGT et al. 1997). It is known from scattered localities on rhe Adanric versant in central Veracruz (PEREZ-HIGAREDA & SMITH 1991), Oaxaca (CASAS-ANDREU et al. 1996), Chiapas, Ta basco, southwestern Campeche and southern Quintana Roo, and on che Pacific versant from Jalisco (CASAS-ANDREU 1982, Ri\MTREZ-BAUTISTA 1994), Colima (SMITH & PEREZ-HIGAREDA 1989), Guerrero (DAVIS & DocoN 1959) and Chiapas (ALVAREZ DEL TORO 1973, 1982, JOHNSON 1989). Natural history: This large, active snake inhabits tropical deci duous forest, evergreen seasonal forest and rain forest. Ir is prima rily terrestrial and nocturnal, bur can also be found active dur ing rhe day. It usually forages at night iJ1 primary or secondary forest, often along screams. Ir feeds mostly on snakes, including nauyacas (Bothrops aspe1) and other venomous snakes, which some times may be about as long as the snake eater itself (CAMPOS V1LLANUEVA & CABRERA-GUZMAN 2009). Occasionally frogs, lizards and mammals are also eaten (RAMIREZ-BAUTISTA 1994, SoL6RZANO 2004). CLelid scytalina is oviparous.
Genus Coniophanes
HALLOWELL, 1860
This genus contains approximately 18 species of small to mode rate-sized snakes which arc rerrestrial and primarily nocrurnal. The overall range of the genus extends from southern Texas and Sinaloa soum mrough Ceacral America and western South Ame rica into northwestern Peru (BAILEY 1939, SMITH & TAYLOR 1945, PETERS & OREJAS-MJRANDA 1970). Thirteen species are found in Mexico. The rail of these snakes is easily broken (unspecialized pseudaucoromy) and about 40% of adults in most samples have blunt tails caused by partial rail loss (MENDELSON 1992, SAVAGE 2002). The posterior teeth on the maxillary are enlarged, grooved and associated with venom glands. Although mese snakes are basi cally harmless co humans and usually make no attempt co bite, their toxic secretions may cause local pain with numbness, swelling and bleeding. Other features char characterize che genus are body cylindrical and moderately slender; tail medium in length; head moderately distinct from the neck, rounded or somewhat pointed in dorsal view; eyes small co moderate and with a round pupil; basic colubrid complement of enlarged head places; dorsal scales
210
Species Accounrs: Alethinophidia: Dipsadidae
Fig. 256: Coniophanes alvarezi, Comitan, Mescta Cenrral de Chiapas, elevation 1,950 m. A. RAMIREZ VEL\ZQUEZ
Fi g. 257: Coniophanes bipzmctatus, Carrecera Las Choapas-Cuichapa, Veracruz. L. CANSECO-MARQUEZ
Fig. 258: Coniophanes bip1mctatus, venrral view, Lago de Catemaco, Sierra de Los Tuxtlas, Veracruz, elevation 400 m. P. HEIMES
Species Accounts: Alerhinophidia: Dipsadid ae
211
Fig. 259: Coniophanes fissidens,
La Victoria, Sierra de Los Tuxtlas, Veracruz, elevation 400 m.
P.
HEIMES
Fig. 260:
Coniophanes imperialis imperialis,
Tenampulco, Puebla.
L. CANSECO-MARQUEZ
Fig. 261: Coniophanes imperialis clavatus,
Tuxda Gutierrez, Chiapas, elcvacion 510 m.
A.
RAMIREZ VELAZQUEZ
21'2
Species Accounts: Alechinopbidia: Dipsadidac
smooth and in r7 co 25 rows at midbody, with a reduction ancerior co the vent; subcaudals and anal place divided.
Coniophanes alvarezi CAMPBELL, 1989 Chiapan srripeless snake; cafetalera lisa de Chiapas
Key to the Mexican Species of the Genus Coniophanes 1a. Dorsal scale rows at midbody 23 tO 25 ............................... 2 1b. Dorsal scale rows at midbody 21 or fewer ........................... 5 2a. Dark lateral stripes 4-6 scales wide ... Coniophanes piceivittis 26. Dark lateral stripes nor more than 3 scales wide ................ 3 3a. Dark vertebral srripe about 4 scales wide .............................. ................................................. Coniophanes michoacanensis 36. Dark vertebral stripe 6-8 scales wide ................................. 4 4a. Dark lateral stripes 0.5-1.5 scales wide, lower edge color grading into ventral coloration ........... Coniophanes schmidti 46. Dark lateral stripes 2-3 scales wide, lower edge color sharply contrasting from color of veneer ............ Coniophanes taylori 5a. Veneer with a double row of large, regular dark spots, one pair on each ventral; chin moccled .................................... 6 5b. Veneer immaculate or with numerous small dark spocs; chin usually sparred, bur not mottled (unless head is black) ..... 7 6a. Ventrals fewer chan 150; no conspicuous lateral dark stripe ... .................................................. .... Coniophanes bipunctatus 6b. Ventrals more than 150; conspicuous lateral dark stripe present ......................................... Coniophanes qieinquevittatus 7a. Head and neck black; body unicolor orange or rusty red, rarely striped .......................................................................... 8 7b. Head and neck not black ................................................. JO
Coniophanes alvarezi CAMPBELL 1989: ro36. Type locality: "n.3 km ESE Teopisca, 2073 m elevation, Chiapas, Mexico". Identification: A smail snake with a uniform brown dorsum and yellow veneer; 19 dorsal scale rows ar midbody. Description: TL up to 52 cm. The dorsal coloration is uniform brown or olive brown, the ventral surfaces are immaculate pale yellow, except for the throat and anterior pan of the body where a small amounr of fine black peppering is present. A black 1 ine runs along tbe sides of che head, extending from che tip of the snout to the angle of che mouth and separating the white supra labials from the brownish coloration above; che supralabials are speckled with black. The iris is copper-colored, with heavy SLLf fusion of black on the lower half. Juveniles have a dark brown head cbat contrasts wirh the pale brown of the dorsum, and very fainc dark lines on the vertebral and fomch scale rows. There are 7 supralabials, 9 infralabials, 1 loreal, 1 preocular, 2 poscoculars, 1+2 temporals, 134-143 ventrals and 54-64 divided subcaudals. Distribution: Coniophanes afvarezi is known only from che Meseca Cencral of Chiapas at elevations from 2,012 to 2,134 m.
Natural history: All specimens of rhe type series were found in temperate mesic pine-oak forest under rocks or logs at che edges of cleari11gs (CAMPBELL 1989).
Coniophanes bipunctatus
(GUNTHER,
1858)
Two-spotted snake; cafecalera de agua, cabaquilla 8a. Head and neck black, followed by a light collar I scale long ......................................................................................... 9 8b. Head and neck black, followed by a light collar about 4 scales long ......................................... Coniophanes meuznocephalus 9a. Dorsum rusty red; light collar usually black-edged posteriorly ..................... ... ..... .... ......................... Coniophanes uzteritius 96. Dorsum orange red; light collar with only a slight amount of dark pigmenc posteriorly ......................... Coniophanes sarae IOa. Dorsal scale rows at midbody 17 ..... Coniophanes meridanus 10b. Dorsal scale rows at midbody 19 or 21 ............................. II na. Dorsum unicolor brown withour longitudinal stripes .......... ...... ....... ............................ ... .. ... .......... Coniophanes alvarezi ub. Dorsum with longitudinal stripes .................................... 12 12a. Pale, dark-bordered stripe running along side of head from snout rip a.bove eye co posrerior temporal scales ................... ........................................................ Coniophanes imperialis 126. Head markings different, if a dark-bordered stripe is present, ic is limited to che temporal region ...... Coniophanes fissidens
Coronel/a bipunctata GONTHER 1858: 36. Type locality: unknown. SCHMIDT (1941) suggested that cl1e type specimen probably came from Belize. Glaphyrophis pictus ]AN 18636: 304. Coniophanes bipzmctatu.s CoPE 1866a: 128 . Coniophanes bipunctatus biseriatus SMITH 1940: 59. Type locality: Palenque, Chiapas, Mexico. Identification: A moderate-sized brown snake; dorsal pattern of suipes usually indistinct, no conspicuous lateral dark stripe; belly with a regular double row of black spots, one pair on each vent ral scure; ventrals fewer than 150; usually 21 dorsal scale rows ac midbody. Description: TL up co 87 cm. The dorsum is reddish brown, dark brown or medium brown and usually has a poorly developed striped pattern. The 3-5 middorsal scale rows are darker than che more lateral rows and usually are marked wich an indistinct black vertebral line. Laterally, a dark brown stripe is present on scale row 4 and the adjacent pardons of rows 3· and 5; th_is stripe is bordered dorsally by a thin black line. The head is dark brown above. Dark vermiform markings are presenc on the white labials, chin and gulars. The belly is cream or pale orange and is marked with a double series of large black spots wltich are arranged in
Species Accouncs: Alethinophidia: Dipsadidae
pairs, one OD each ventral scale.The spors are largest on the chroar and anrerior part of the body and become smaller posteriorly. There are 7-9 supralabials, 8-ll infralabials, 1 lorea.l, r preocular, 2 poscoculars, r+1 1 r+2 or 1+3 remporals, 124-147 ventrals and 72roo divided subcaudals. Distribution: Coniophanes bipunctatus ranges ac low elevations (generally less rhan 500 m) on rhe Aclanric versant from south easrern San Luis Potosi (vicinity ofTamamnchale) and northern Puebla sourhward through Veracru2, norrheascern Oaxaca, Ta basco, northeastern Chiapas and coasral areas of the Yucatan Peninsula to norcl1western Panama (SMITH 1943a, SMITH & TAYLOR 1945, PEREZ-HIGAREDA & SMITH 1991, LEE 1996, VoGT et al. l997). SMJTH (r940) recognized rwo subspecies on the basis of color parrern and number of subcaudal scales. He assigned specimens from Veracruz, wesrern Campeche and norili-ce.ntral Ch iapas to the subspecies C. b. biseriatus and referred all ocher populations ro ilie nominate subspecies, C. b. bipzmctatus. Co N,INT (19656) and WILSON & MEYER (1985) quesrioned the valid ity of iliis arrangement and pointed our rhar variation in chis species is poorly understood. Natural history: This terresuial and semi-aquatic snake inhabits marshes and swamps in rropical deciduous foresr, evergreen sea sonal forest and rain forest. CAMPBELL (1998a) found individuals ar nighr lying on aquaric vegerarion and by day under logs or debris ar the edges of bodies of water. Coniophanes bipunctatus feeds on frogs, eels and probably small lizards (SOLORZANO 2004). Up to 6 eggs are laid during the rainy season. SMITH (1940) found a clutch of 5 eggs in a rorren scump on the bank of a stream in Chiapas. Coniophanes fissidens (GUNTHER, 1858) Yellow-bellied snake; culebra panza amarilla Coronellafissidens GONTHER 1858: 36.Type localiry: Mexico and America; restricted to San AndresTuxrla, Veracruz, Mexico, by SMITH & L\YLOR (1950b). Coniophanesfosidens Co PE 1860a: 248. Coniophanes punctigularis COPE 1860a: 248. Type localiry: Hon duras. C. [oniophanes] proterops Cof'E 1860a: 249. Type locaJjry: Vicinity of Jalapa, Veracruz, Mexico. Coniophanes fissidens fosidens BAILEY 1939: 14. Coniophanes fissidens proterops SMITH 19416: 105. Coniophanes fosidens di.sperms SMITH 19416: 106. Type locality: El Limoncito, Guerrero, Mexico. Coniophanes fissidens punctigularis SMJTH 19416: 107. Co11iophanes fissidens conve1gens SHANNON & SMITH 1950: 499. Type locality: 6 mi. NW Tihuarl:in, Veracruz, Mexico. Identification: A small brown, indisr.incrly striped snake; belly sprinkled with small black spots; no light temporal stripe through cop of orbit, dorsals in 19-21 rows at midbody.
213
Description: This species rarely exceeds 55 cm inTL, bur occa sionally may reach 80 cm. The dorsal colorarion is brown. The scales of rhe vertebral row may be bordered wiili black, creating a poorly developed middorsal line. A thin black dorsolateral stripe on scale row 5 which usually is bordered above by a light stippled line separates the pale brown color of rhe middorsum from the darker brown color on the sides.The dorsolateral light line extends forward to the upper side of ilie neck where it usually rerminaces in a pair of black-edged, cream spots; a few black-edged light spots are usually presenc on rhe upper temporal region, and a white narrow stripe exrends across rhe supralabials onto the side of the neck. The belly is cream or pale yellow, sometimes with a pinkish tinge, and is marked with numerous dark spots which ofren are more or less arranged in rwo lines on the outer edges of the venc rals.The chin and iliroat are densely stippled with black pigment. There are 8 supralabials, 8-10 infralabials, r loreal, r or 2 preocu lars, 2 posroculars, 1+2 temporals, 109-146 ventrals and 57-w3 divided subcaudals. Distribution: Coniophanesfissidens ranges at low and moderate elevations (up co 1,400 m) on rhe Arlancic versant fromTamau lipas, and on the Pacific versanc from Mkhoacan, southward through Central America (excluding the outer Yucatan Penin sula) to Colombia and Ecuador (KOHLER 2003). FISHER (1968), as well as oilier authors, recognized six subspecies, five of which are found in Mexico: On the Aclanr.ic versant, C. f convergens occurs from southernTamaulipas through southeastern San Luis Potosi, noriliern Queretaro, eastern Hidalgo, noriliern Puebla and noriliern Veracruz; C. f proterops in central Veracruz; and C. f fissidens from northeastern Oaxaca through southern Veracruz, northern Chiapas, and southern Tabasco to noriliern Central America. On the Pacific versanr, C. f di.sperms ranges from sourh wesrern Michoac:in southward to west of rhe Isthmus ofTehuan tepec, and C. f punctigularis from easr of the Isthmus through coastal Chiapas sourhward to Honduras. Distribution informa tion for Mexico is given in BAILEY 1939, SMITH 1943a, SMITH & TAYLOR 1945, DUELLMAN 1961, 19656, ALVAREZ DELTORO 1973, 1982, SALDANA DE LA RrvA & PEREZ RAMOS 1987,JOHNSON 1989, PEREZ-HIGAREDA & SMITH 1991, CASAS-ANDREU et a!. 1996, LEE 1996, BENITEZ GALVEZ 1997, CANSECO-MARQUEZ & GUTIERREZ MAYEN 20066, GUTIERREZ-MAYEN & SALAZAR ARENAS 2006, FARR et al. 2009, DrxoN & LEMOS-ESPINAL 2010 and LEMOS ESPINAL & DJXON 2013.
Natural history: This terrestrial snake inhabits tropical deciduous forest, evergreen seasonal forest, rain forest and cloud forest. le forages in rhe leaf licrer of the foresr floor by day and at nighr. Ir feeds mostly on frogs of rhe genus Cra11gastor, but a wide variety of prey items is also earen, including ocher anurans, salamanders, lizards (anoles and skinks), small snakes (including individuals of irs own species) and rhe eggs of amphibians and reptiles (MINTON & SMITH 1960, LANDY et al. 1966, SEID 1985). Juveniles also con sume inverrebraces (SAVAGE 2002). Clurches ofJ-8 (usually 2-4) eggs are laid between March and July (ZuG et al. 1979, UMP BELL 1998a, SoL6RZANO 2004).
214
Species Accounts: Alethinophidia: Dipsadidae
Fig.
262:
Coniophane, lateritius,
Minaticlan, Colima.
C.
GRUNWALD &
J. JONES
Fig. 263:
Coniophanes lateritius,
striped morph, Colima.
C.
GRUNWALD & J. JONES
Fig.
264:
Coniophanes meumocephalus,
Tepozrlan, Morelos.
P.
HEIMES
Species Accounts: Alethinophidia: Dipsadidae
215
Fig. 265: Coniophanes meridanus,
Yucatan Peninsula.
J.
ORTIZ
Fig. 266: Coniophanes piceivittis,
Tuxtla Gurierre2, Chiapas, elevation 510 m. A. RAMIREZ YEu\ZQUEZ
Fig. 267: Coniophanes schmidti,
Calakmul, Campeche. L. CANSECO-MARQUEZ
216
Species Accounts: Alerhinophidia: Oipsadidae
Coniophanes imperialis (KENNICOTT, 1859) Regal black-striped snake; culebra de raya negra, viencre rojo, yak' ik'kum (Lacand6n Maya) Taeniophis imperialis KENNICOTT [in BAJRD] 1859: 23. Typ e locali ty: Matamoros, Tamaulipas, Mexico. Glaphyrophis lateralis J AN 1863b: 304. Dromicus clavatus PETERS 1864: 388. Typ e localicy: Mexico; restric ted co Porrero V iejo, Veracruz, Mexico, by SMITH & TAYLOR (1950b). Coniophanes imperalis imperalis COPE r900: 1090. Coniophanes imperialis copei H ARTWEG & OuvER 1938: 4. Type locality: between Cerro Guengola and Tehuantepec, Oaxaca, Mexico. Coniophanes imperialis clavatus BAILEY 1939: 35. Identification: A small brownish snake with a pattern of dorsal and lateral stripes, dorsal stripe often indistinct; light temporal stripe through top of eye; dorsal scales in 19 (rarely 21) rows at midbody. Description: This relatively slender snake reaches up co 60 cm
TL. The dorsal ground color is brown; the can or medium brown middorsal area is separated from the dark brown sides of the body by a light dorsolaceral scripe which often is indistinct. A black vertebral line, which may be solid or broken inco a series of dashes, usually runs the length of the body. The cop of the head is medium brown or dark brown. A white or cream stripe runs from the tip of the snout through the upper edge of the eye and onto rhe temporal region; ic may be continuous with the light dorsolareral stripe on rhe body, but more often terminates in the temporal region, in which case there is a distinct white nape spot be.tween the posterior end of the head stripe and the anterior end of the body stripe; sometimes rhe nape spot is fused with the body stripe. The labials, chin and anterior part of the belly are white, grading to yellow or yellow-orange at midbody and orange or salmon near the vent. There are dark punctuations on the chin, and a series of irregular black midventral spots are usually con centrated on the anterior part of the belly. There are 8 supralabials, 9-10 infralabials, I loreal, I preocular, 2 posroculars, 1+2 temporals, n4-14r ventrals and 62-94 divided subcaudals. Distribution: Coniophanes imperialis ranges on the Atlantic ver sanr from southern Texas through eastern Me,_ico (including the Yucatan Peninsula) to northern Honduras. Ir also occurs locally on the Pacific versanc in che Isthmus region of southeastern Oaxaca. BAILEY (1939) recognized three subspecies. The nomi nate subspecies is found from Texas southward co northern Veracruz, and C i. clavatus ranges from central Veracruz co Hon duras (BAILEY 1939, SMITH 1943a, SMITH &TAYLOR 1945, TAYL OR 1953, M ARTIN 1958, P.EREZ-HIGARED A & SMITH 1991, LEE 1996). Specimens from southern Oaxaca have been allocated co C. i. copei (SMJTH &TAYLOR 1945, CASAS-ANDREU et al. 1996). Conio phanes imperialis is commonly found ar low and moderate eleva tions, but in Chiapas it was reported from elevations up to 2,000 m OottNSON 1989).
Natural history: Coniophanes imperialis inhabits tropical decidu ous forest, evergreen seasonal forest, rain forest and marginally
pine-oak forest. le occurs in forested areas as well as in open habitats and along the edges of marshes. It is terrestrial and pri marily nocturnal, buc occasionally individuals are seen by day crawling in the leaf litter in search for prey. Ochers have been found beneath ronen logs and other surface debris (HENDERSON 1976a, LEE 1996). The diet consists mosrly of frogs and lizards, but small snakes and insects are also eaten (HENDERSON & HoE VERS 1977b, CAMPBELL 1998a). Females lay 3-ro eggs between May and August, and the young hatch after approximately 40 days; neonates measure about 150 mm in TL (f\.LVAREZ DEL TORO 1973, 1982, CAMPBELL 1998a). This snake is generally inoffensive and rarely bites. BR OWN (1939) reported that a bite caused burn ing, itching, swelling and red discoloration in che region of the fang punctures, and that these symproms lasted for two to three days. LEE (1996) reported experiencing incense throbbing pain and muscular stiffness char persisted for several days after a bire of a small specimen. Coniophanes !ataitius COPE, 1862 Srripeless snake; culebra lisa Coniophanes !11teritius COPE 1862 ("r86r"): 524. Type locality: Guadalajara, Jalisco, Mexico. Tachymenis lateritia G ARMAN 1884a: 61. Erythrofampms !ateritius COPE 1887: 78. Coniophanes I. lateritius WELLM AN 1959: 127. Identification: A small snake with a ruscy-red dorsum; dorsal sm faces of head and neck black, followed by a light collar one scale long and usually black-edged posteriorly; black nuchal hood nor extending to ventrals; dorsal scales in 19 rows at midbody. Description: TL up to about 60 cm; apparencly females are larger rhan males. The dorsal coloration is bright red or brick-red, rhe middorsum is somewhat obscured. In some individuals, the pos terior two-thirds of the body and the entire dorsal surface of the rail are nearly black; ocher individuals may have five pale yellow stripes running along the body and rail. The top of rhe head and neck are black, rhe black hood extending posterior co che pari etals a length of 6-8 scales on the vertebral scale row. The black coloration extends on each side co scale row 2 or 3. The black nuchal hood is bordered posteriorly by a pale yellow collar one scale long which in turn is edged posteriorly with black pigment. There are numerous Jjghr dots on the top and sides of the head. The ventral surfaces are cream-colored, chin and throat are marked with numerous black spots. The iris is dark brown. There usually are 7 supralabials, 9 infralabials, I loreal (rarely absent), 1 preocu lar, 2 posroculars, 1+2 temporals, 137-154 ventrals and 82-99 di vided subcaudals. Distribution: Pacific versanc from central Sinaloa southward to central Guerrero (BA1l.EY 1939, SMITH & GRANT 1958, WELLMAN 1959, TANNER & R OBISON 1960a, McDrARMlD 1963, PoNCE CAMPOS & SMITH 2001, C. GR0N\xrALD, pers. comm.). Conio phanes lateritius was also listed for Sonora (AMBfA MouN A 1969), bur no localities were given. The vertical distribution ranges from near sea level to about 1,600 m (P ONCE -CAMPOS & SMITH 2001).
Species Accounrs: Alechinophidia: Dipsadidae
217
Natural history: This uncommon terresrrial snake inhabirs rrop ical scrub forest, semi-deciduous forest and pine-oak forest.
lateral stripes cover rows 3 and 4. The top of the head is dark brown. A light temporal srripe runs from the rip of the snour through the upper edge of the eye and onto the temporal region. Mose specimens have a lighr nuchal collar bordered with black; Coniophanes melanocephalus (PETERS, 1870) the collar is interrupted middorsaUy by dark pigment, ventrally it is continuous with the light ventral coloration. In some speci mens the collar is reduced to a pair of light nuchaJ spots. The Black-headed srripeless snake; culebra lisa de cabeza negra labials and ventral surfaces are light can or reddish tan with a few Tachymenis melanocephala PETERS 1870: 874. Type locality: "Pueb- dark spots anteriorly. There are usually 7 supralabials, 1 loreal, la", restricted to Matamoras [Izucar de Matamoros], Puebla, r preocular, 2 postoculars, 1+2 temporals, 122-134 venrrals and Mexico, by SMITH & T.wLOR (19506). 83-87 divided subcaudals. Erythrolamprus melttnocepha/11s CoPE 1887: 78. Coniophanes !ateritius melanocephalus SMITH & GR.ANT 1958: 22. Distribution: Yucatan Peninsula. Coniophanes meridanus is known Coniophanesmelanocephalus PONCE-Cu.ir>os & SMITH 2.001: IO. from several localities in che scares ofYucacan and Quintana Roo, and there are records from adjacent areas of northern Campeche ldentification: A smaJJ snake with a rusty-red dorsum; dorsal sur (McCoY 1969, LEE 1996, CHARRUAU & CEDENO-VAZQUEZ 2005). faces of head and neck black, foUowed by a Light collar about 4 scales long; black nuchal hood extending ro ventrals; dorsal scales Natural history: This is a terrestrial and nocturnal inhabitant of tropical thorn forest and tropical deciduous foresr. in 19 rows at midbody. Description: Adults are commonly 35-50 cm long. The dorsal ground color is bright red or brick-red, the middorsum is some what obscured.The rop of the head and neck are black, tbe black hood extending posterior co rhe parierals a length of 6-ro scales on rhe verrebral scale row; on the neck, the black coloration ex tends lareraJly to rhe ventrals. The black nuchal hood is bordered posteriorly by a pale yellow collar about 4 scales long; only a small amount of dark pigment borders the collar posteriorly. The labi als have some light spots, the chin is cream with black pigment, and the belly is pale yellow and immaculate. The iris is dark red. A single specimen was reported ro have 7 supcalabials, 9 infrala bials, 1 loreal, 1 preocular, 2 posroculars, 1+2 temporals, 156 vent rals and 86 divided subcaudals. Distribution: Rio Balsas Basin of central Mexico. Coniophanes mclanocephalus ranges at 900-1,700 m elevation through much of the Rfo Balsas Depression from southern Michoadn and northern Guerrero ro southwestern Puebla (GARCIA-VAZQUEZ et al. 2006). Natural history: This uncommon terrestrial snake has been found
in tropical semi-deciduous forest and pine-oak foresr.
Coniophanes meridanus SCHMIDT & ANDREWS, 1936 Peninsular srripeless snake; culebra lisa de Yucatan Coniophanes meridanus SCHMIDT & ANDREWS 1936: 179. Type locality: Merida, Yucatan, Mexico. Identification: A smaU reddish brown snake with or without in d istincc dark stripes on anrerior parr of body; a light temporal stripe through top of eye; a pair of light nuchal spots, or a d o r sally interrupted, light collar; dorsals i n 17 rows ar midbody. Description: TL up to about 50 cm. The dorsal ground color usually is reddish brown. There may be very slight traces of dark stripes anceriorly, especially in younger specimens; if present, the middorsal stripe is confined co the verrebral scale row and the
Coniophanes michoacanensis FLORES-VILLELA & SMITH, 2009 Michoacan black-striped snake; culebra rayada de Michoacan Coniophanes piceivittis VARGAS-SANTAMARiA & FLOREs-VtLLEL.A 2006: 110. Coniophanes michoacanensis FLORES-VTLLELA & Sl'vUTH 2009: 404. Type locality: "El Farito, 8 km NW Caleta de Campos, 17 m elevation, Municipio de Aquila, Michoacan, Mexico (18.1014°N, 102.8173"W)". Identification: A small to moderate-sized snake with three dark stripes; vertebral stripe about four scales wide; lateral stripes about two scales wide, sharply defined on lower edge; dorsal scales in 25 rows at midbody. Description: The holotype, an adult female, measures 56.1 cm in TL. The ground color is pale brown, turning co white-cream on the head and ventrally. The dorsal pattern consists of three dark brown stripes chat originate on the snout, broaden on top of the head and on che cemporal region, and run posteriorly the length of che body and tail. The vertebral dark stripe is tluee and rwo half scales wide, and the lateral dark stripes are about one and rwo halfscales wide almost throughout the length of che body. All stripes contrast sharply with the ground color. The dorsal and lateral dark stripes are separated by an intervening space of 3 4scales of cream ground color. The parietals have small pale docs mediaUy, and each supralabial scale has dark pigment distributed on the lower half. The chin and gular regions are peppered with dark pigment, che remaining ventrals and the subcaudals are im maculate. There are 8 supralabials, 9 infralabials, 1 loreal, 2 pre oculars, 2 poscoculars, 1+2 temporals, 164 ventrals and 87 divided subcaudals. Distribution: Western Mexico. Coniophanes michoacanensis is
known from Pacific coastal areas (up to 800 m) of Michoacan and adjacent northwestern Guerrero (FtoREs-VILLELA & SMITH 2009, MERTZ et al. 2012.).
218
Fig. 268: Conophis lineatus, Yucatan PeninSU Ia.
Species Accounts: Alerhinophidia·· 01· psad"d I ae
].
ORTIZ
Species Accounts: Nechinophidia: Oipsadidae
Fig.
270:
Conophis lineatm, Cacemaco, Sierra de Los Tuxdas, Veracruz, elevation
Fig. 271: Conophis lineatus, Ciudad Cardel, Veracruz.
2.50
m.
2.19
A. RAMiREz VEw\ZQUEZ
P. HEIMES
220
Species Accounts: Akrhinophidia: Dipsadidae
Natural history: A terrestrial and noccurnal inhabitant of tropi cal deciduous forest. The type locali ty has some remnants of pri mary forest surrounded by fruit plantations (FLORES-VILLELA & SMITH 2009).
Coniophanes piceivittis COPE, 1869 Cape's black-striped snake; culebra rayada, tzinkan (Lacandon Maya)
Coniophanes piceivittis COPE 1869b: r49. Type locality: Chihui tan, Oaxaca, Mexico. Coniophanes piceivittis frangivirgatus PETERS 1950: 279. Type lo cali ty : 0.5 mi. E Plan de! Rio, Veracruz, Mexico. Identification: A small, distinctly striped snake; three wide dark stripes separated by two narrow light stripes; vertebral stripe 5-8 scales wide; lateral stripe 3-6 scales wide, usually sharply defined on lower edge; dorsal scales in 23-25 rows at midbody. Description: This relatively slender snake reaches a maximum TL of 57 cm. The dorsal color pattern is composed of three wide dark brown co black stripes, one median and one lateral on each side, which are separated by nvo narrow, cream or can dorsolate ral stripes. The median dark stripe occupies 5-8, the lateral dark stripes 3-6, and the pale stripes in benveen one co one and cwo half scale rows. The pale stripes originate on each side of the rost rum, pass through the upper edge of che eye and extend co the tip of che rail. They may be interrupted in the temporal region, in which case there is usually a nape spot benveen the rwo ends of the srripe. The lower surfaces of the body are cream, tan or orange; labials and chin are sparred with dark brown or black. The nvo subspecies are mainly distinguished by che presence (in C. p. frangivirgatus) or absence (in C. p. piceivittis) of a nape spot. There are 7-8 supralabials, 9-10 infralabials, I loreal, 2 (rarely r) preoculars, 2 poscoculars, 1+2 temporals, 153-171 ventrals and 7694 divided subcaudals; anal ridges are present in adult males.
Distribution: Coniophanes piceivittis occurs ar low and moderate elevations (sea level co 1,400 rn) on the Arlamic versanc from southern TamauJipas southward to ce.nrral Chiapas, and on the Pacific versam from southern Oaxaca co northern Costa Rica (FLORES-VILLELA & SMITH 2009). Two subspecies are recognized, with C. p. frangivirgatus occuring on the Atlantic versanc (MARTIN 1958, PER.EZ-HIGAREDA & SM1TH 1991, BENITEZ GALVEZ 1997, DIXON & LEMOs-EsPJNAL 2010, R,\MfREz-BAUTISTA et al. 2010), and C. p. piceivittis on the Pacific versanc (AtvAREZ DEL ToRo 1973, 1982, C AS AS-ANDREU et al. 1996, 2004). Natural history: This cerresrrial snake inhabits tropical thorn forest, deciduous forest and evergreen seasonal forest. It is cre puscular and nocturnal and has been found in vegetation or leaf litter as well as under rocks and logs. Ir ears small frogs, lizards and occasionally the eggs of other reptiles. Clutches of up co six eggs are laid in July; neonates have been recorded in September and October (S oL6RZANO 2004).
Coniophanes quinquevittatus (DuMERIL, BrnRoN & DuMERIL, 1854) Five-striped snake; culebra de cinco lineas
Homalopsis quinque-vittatus DuMERIL, BIBRON & DuMERIL 1854: 975. Type locali ty: unknown. SMlTH &TAYLOR (r95ob) desig nated El Peten, Guatemala, as the type locality. Calopisma quinquevittatum mexicana ]AN 1863a: 55. Type locality: "Mexico", restricted co Puerco Mexico, Veracruz, Mexico, by SMITH & TAYLOR (19506). Hydrops lubricw COPE 1871: 217. Type locali ty: "Coaczacoalcos River, Veracruz", resrricred co Puerto Mexico, Veracruz, Mexico, by SMITH & TAYLOR (19506). Coniophanes quinquevittatus BAILEY 1939: 26. Identification: A small co moderate-sized, striped snake; 3 dark stripes separated by narrower pale stripes; venter pale with a re gular double row of large dark spots, one pair on each ventral scute; ventrals more than 150; 21 (rarely 19) dorsal scale rows at midbody. Description: TL up co 87 cm. The dorsal color pattern consists of a broad dark brown median stripe and a dark slate-gray or gray-brown stripe on each side; these dark stripes are separated by cream or pale brown dorsolateral stripes. The median srripe is usually marked with an indistinct black vertebral line. The dark lateral stripes originate on the snout, pass through the eye and across che temporal region and occupy scale rows 3 co 5 and adja cent portions of rows 2 and 6 at midbody. The top of the head is brown and heavily pigmented with dark gray. The lower sides of the body are white, cream or pale yellow. Tbe belly is light tan co cream anteriorly, usually becoming yellowish or pinkish pos teriorly. Labials and chjn are boldly spotted wirh dark brown or black, and there is a pair of large black spots on each ventral scale. These spars may coalesce co form a zigzag line on the under surface of the rail. Often there are smaller spots on the midvem ral area. There are 8 supralabials, 10-11 infralabials, I loreal, 1 pre ocular, 2 poscoculars, r+2 temporals, 157-164 ventrals and 63-70 divided subcaudals; anal ridges are present in adulc males. Distribution: Coniophanes quinquevittatus occurs at low eleva tions (up co 200 m) on che Atlantic versant of southern Mexico and northern Central America. This species has a highly disjunct distribution in the lowlands of southern Veracruz (between Alva rado and Coatzacoalcos), che northwestern coastal area of the Yucatan PeninsuJa, and rhe central Pecen of northern Guatemala (C AMPBELL 1998a, PE REz-HrG AREDA & SMITH 1991, LEE 1996, VOGT er al. 1997). Natural history: Coniophanes quinquevittatus occurs in tropical thorn foresr, deciduous forest and evergreen seasonal forest. This semi-aquatic species is closely associated wirh marshy or swampy situations with thick vegetation along the edges of lakes and rivers. Ir is nocturnal and feeds on frogs and fish (CAMPBELL 1998a).
Species Acc�uncs: Alethinophidia: Dipsadidae Coniophanes sarae Por,;cE-CAMPOs & SMITH, 2001 Sara's srripeless snake; culebra lisa de Sara Coniophanes sarae PONCE-CAMPOS & SMITH 2001: 10. Type loca lity: "near Tehuamepec, municipality ofChinicuila ( 1 8°42-'oi'N, Io3°18'22.3"W )1 1 390 m, Michoacan", Mexico. Identification: A smalJ snake with an orange-red dorsum; dorsal surfaces of head and neck black, followed by a light collar one scale long; black nuchal hood not extending to ventrals; dorsal scales in 19 rows ar midbody. Description: Coniophanes sarae is known only from two speci mens (PoNCE-CANrPos & SMITH 2001). Adults probably reach about 50 cm in TL. The dorsal colorarion is orange-red.The top of the head and neck are black, the black hood extendi11g poste rior to the parietals a length of 5 scales on the vertebral scale row. The black coloration ends on each side at the second scale row. The black nuchal hood is bordered posteriorly by a pale yellow collar one scale long.There are numerous light dots on the rop and sides of the head. An irregular, discontinuous light line passes through the supralabials, curving down to the ventrals and joining with the ventral end of the light nuchal collar. The iris is dark brown. There are 7 supralabials, 9 infralabials, r loreal, 2 preocu lars, 2 postoculars, 1+2 temporals, 151-156 ventrals and 85-91 di vided subcaudals. Distribution: Coniophanes sarae is known only from intermediate elevations in the Sierra de Coalcoman of southwestern Micho acin.The relationship berween chis species and C. melanocephafus in rhe Rio Balsas/RioTepalcarepec Depression remains co be cleared. Furthermore, ir is unclear whether individuals from coastal areas of Michoacin are of this species or C. lateritius. Natural history: Coniophanes sarae is an in habitat of cropical semi-deciduous forest. Coniophanes schmidti BAILEY, 1937 Schrnjdt's black-striped snake; culebra rayada de Yucatan Coniophanes schmidti BAILEY 1937a: 1. Type locality: Chichen Itza, Yucatan, Mexico. Coniophanes piceivitti.s schmidti NEILL & ALLEN 1960: 1 45. Identification: A small co moderate-sized snake with a broad dru·k middorsal stripe bordered by narrow, black-edged, light dorso laceral scripes; dorsals in 23-27 rows at midbody. Description: TL up to 70 cm. A broad, dark brown to almost black middorsal stripe is bordered on each side by a distincrive narrow cream or yellow dorsolateral stripe on scale rows 8-9. The pale dorsolateral stripe, which is black-edged vencrally, may begin on the anterior part of the body or may be continuous with a narrow, dark-edged, cream or yellow head stripe that originates on the rostrum and passes through che upper edge of the orbit and upper temporal region. In case char the stripe is discontinuous,
221
it may form a discrete light nape spot between its two ends. The sides of the body (scale rows r-6 and the lower pare of 7) are pale yellow to pale brown. The top and sides of the head are reddish brown. The ventral surfaces and labials are cream or light tan; labials and gular region usually are peppered with a little dark pigment. There usualJy are 8 supralabials, 10 infralabials, I loreal, 2 (rarely r) preoculars, 2 poscoculars, 1+2 temporals, 161-170 vent rals and 91-104 divided subcaudals; anal ridges are present in adulr males. Distribution: Coniophanes schmidti occurs throughout the Yuca tfo Peninsula southward co cl1e Peten region in northern Guate mala. The vertical distribution ranges from near sea level co about 300 m (MAHRDT 1 969, McCOY 1969, LEE 1996, CAMPBELL 1998a, FERREIRA-GARCIA & CANSECO-MARQUEZ 2006). Some authors (e.g., WERLER & SMITH 1952, WILSON & MEYER 1985) have con sidered C. schmidti a subspecies of Coniophanes piceivittis. Natural history: This terrestrial and nocturnal snake inhabits tro
pical thorn forest, tropical deciduous forest and evergreen sea sonal forest. Ir feeds on frogs and lizards. 4 or 5 eggs are laid during the rainy season (CAMPBELL 1998a). Coniophanes taylori HALL, 1951 Taylor's black-striped snake; culebra rayada de Taylor Coniophanes piceivittis taylori HALL 1951 : 202. Type locality: Agua de Obispo, Guerrero, Mexico. Coniophanes tayfori FLORES-VILLELA & SMITH 2009: 404. Identification: A smalJ snake with three black stripes; vertebral stripe about six. scales wide; lateral stripes about 3 scales wide, sharply defined on lower edge; dorsal scales in 25 rows at mid body.
Description: TL up co about 50 cm. The dorsal pattern consists of three black stripes on a ground color of cream, the stripes commencing on the snout and running the length of the body and tail.The vertebral black stripe occupies about 6 scale rows and the lateral stripes 2-3 scale rows. All stripes contrast sharply with the ground color. The dorsal and lateral dark stripes are se parated by ru1 intervening space of 2-3 scale rows of cream ground color. Anteriorly, the median black stripe becomes brown on top of the head, with rhe parietals having a small pale ocellus medi ally. There are blackish spots on the chin and infralabials, the venter is immaculate.T here are 8 supralabials, 9-10 infralabials, r loreal, 2 preoculars, 2 postoculars, 1+2 temporals, 1 64-173 vent rn.ls and 82-90 subcaudals; anal ridges are pL'esent in adult males. Distribution: Coniophanes taylori is known only from rhe Aca puko-Chilpancingo area in central Guerrero, from near sea level to about 1,400 m elevation (HALL 1 951, DrxoN et al. 1962, SAL DANA DE LA RrvA & PiR£z RAMOS 1 987, SANCHEZ & L6rEz FoRMENT 1988, FwREs-VJLLELA & SMITH 2009). Natural history: This cerremial s□ake inhabits tropical decidu ous forest (SALDANA DE LA RrvA & PEREZ RAMOS 1987).
222
Species Accounts: Alethinophidia: Dipsadidae
Fig.
272:
Conophis vittatus,
Sierra de Huautla, Morelos, elevation 950 m. P. HEIMES
Fig. 273:
Conophis vittatus,
Puerto Arista, Tonala, Chiapas, elevation 10 m. A. RAMiREZ VELAZQUEZ
Fig. 274:
Conophis vittattts,
Tuxtla Gutierrez, Chiapas, elevation 510 m. A. RAMiREZ VE1..,\ZQUEZ
Species Accounts: Alethinophidia: Dipsadidae
Fig. 275: Cryophis hallbergi, Sierra Juarez, Oaxaca.
Fig. 276: Habitat of C1yophis hallbergi and Geophis duellmani: Cloud forest near Vista Hermosa, Sierra Juarez, Oaxaca.
223
C. RODRIGUEZ
P. HEIMES
Species Accounts: Alechinophidia: Dipsadidae
224
Genus Conophis
PETERS, 1860
This genus contains two species of moderate-sized, diurnal and terrestrial snakes. They are distributed across the semi-arid and sub-humid coastal plains and foothills from Mexico to Costa Rica (WELLMAN 1963). PtREz-H1GA.R.EDA et al. (2002a) described a new species, Conophis morai, from southern Veracruz, on the basis of a single specimen. It has a color pattern which the authors con sidered unique within the genus. However, the body pattern of the holocype of C. morai seems to be nearly identical to che one seen in other specimens of C. lineattts from southern Veracruz (WILSON & JOHNSON 2010). Therefore I consider C. morai as a synonym of C. lineatus. These snakes are extremely fast and agile and generally are quick to bite. They possess well-developed rear maxillary teeth and fairly potent venom, which in humans causes pain, local sweiling and discoloration. Symptoms may last for rwo or more days, and wounds rend to bleed for an hour or more, suggesting that these snakes may possess an anricoagulanr in their venom (WELLMAN 1963, C AMPBELL 1998a). Orber features thac characterize the genus are body rather srout and cylindrical, rail medium i11 length; head narrow, conical and barely wider than the neck; eyes moderately large and with a round pupil; basic colubrid complement of enlarged head places, including a slightly concave rostral (adapted for burrowing); dorsal scales smooch, in 19 rows at midbody and 17 rows anterior ro the vent; subcaudals and anal place divided. Key to the Species of the Genus Conophis ra. Dorsurn with 4 to 10 black stripes chat are usually less than two scale rows wide; 7 or 8 supralabials ........... Conophis lineatus 16. Dorsum with 3 or 4 black stripes that are often rwo or more scale rows wide; usually 7 supralabials ....... Conophis vittatus Conophis lineatus (DuMERlL, BIBRON & DuMERIL, 1854) Central American road guarder; guarda caminos centroameri cana, sabanera, czinkan (Lacand6n Maya) Tomodon lineatum Dt.JMERJL, BIBRON & DuMERIL 1854: 936. Type locality: "Mexique'', restricted to Veracruz, Veracruz, Mexico, by SMITH &. TAYLOR (19506). Conophis lineatus COPE 1864: 167. Conophis concolor CoPE 1867= 318-319. Type locality: "Yucatan'', restricted to Ch.ichen ltza, Yucatan, Mexico, by SMITH & TAY LOR (19506). Conophis pulcher CoPE r869a: 308. Type locality: "Peren" or "Ve rapaz", Guatemala. Conophis pulcher p!agosus SMITH 1941h: 121. Type locality: Tonala, Chiapas, Mexico. Conophis lineatus lineatus SMITH 1941h: 122. Conophis lineatus dunni SMITH 1942a: 395. Type locality: Mana gua, Nicaragua. Conophis morai PEREZ-H.iGAREDA, L6PEZ-LUNA & SMJTI-1 2002a: 27. Type locality: "southeastern slope of San Marrin Tuxrla Volcano, 1050 m", Veracruz, Mexico.
Identification: A rather srour, striped snake with a poi aced snout; dorsal color pattern consisting of 3-10 narrow black stripes; usu ally some stripes broken into series of dashes; stripes sometimes reduced to series of dots throughout body or only on nape; co loration of areas between stripes variable; first scale row pigmen ted; a pair of diffuse black spots on each ventral presenr or not; usually 8 supralabials; dorsals smooth and in 19 rows at midbody. Description: TL up to u6 cm. The dorsal ground color is pale brown, can, olive, cream, olive gray, bluish gray or white. Three subspecies are mainly distinguished by che number of black stripes extending che length of rhe body. Conophis I. lineatus from the Arlancic versant of Mexico usually have two black lines on each side of the body: a lateral line on scale row 4 and a dorsolareral line on row 7. Both lines initiate on rhe snout, che dorsolateral line extending over the top of the head and nape onto the body, and the lateral line passing th.rough the eye as a broad stripe and nar rowing to a chiJ1 line on che nape ro continue onto the body. Sometimes there is a third line present on the first scale row, bur usually chis row is darkly pigmenrcd witl1out forming a discrete line. There is much variation in rhe colorarion of the areas be tween rhe black lines, but in general the area between rhe lateral and dorsolareral lines is much Lighter than the middorsal and vent rolateral areas, which may result in a contrasting pacrern of broad dark and light stripes. In some specimens secondary dark lines are present in the form of dashes on the posterior pare of the body and rail. On the rail, che secondary and primary lines on adjacent scale rows sometimes fuse into a single broader stripe. Conophis l. concolor from che Yucatan Peninsula is essemially scripeless, except for a broad brown ocular stripe that extends from rhe snout through the eye and onto the sides of the neck. The ocular stripe has narrow black or dark brown borders. On rhe nape, small dark brown dots are often present on che rips of the scales of rows 4 and 7; posteriorly, such docs may be present on row 4 or 7, bur never on both. Conophis L. dunni from the Pacific coastal region of Chiapas has four or five narrow black stripes on each side of the body: (1) on scale row r, (2) on the upper half of rhe second and lower parr of the third rows, (3) on scale row 4, (4) on scale row 7, and (5) a paravercebral stripe on scale row 9. Sometimes the second and third stripes are fused resulting in only 8 stripes. The stripes on the first and third row sometimes are present as disconrinuous rows of dashes. The areas between the stripes usually are pale brown. The supralabials are immaculate white or have dark ventral margins; chin and throat are unmarked or have a few small, scattered black spots. The belly is white or cream, with or wichour black spots on rhe sides of the ventrals. There are 8 (rarely 7) supralabials, 9-ro infralabials, r loreal, 1 preocular, 2 (rarely 3) posroculars, 2+2 (rarely 2+3) tem porals, i55-178 ventrals and 56-80 divided subcaudals. Distribution: On the Atlantic versant from Veracruz, on the Paci fic versant from western Chiapas southward to Cosra Rica. The two versants in Mexico are inhabited by difl:erent subspecies. On the Atlantic side, C. !. lineatus ranges from central Veracruz ro Tabasco (PtREz-H1GAREDA & SMITH 1991, REYNOSo-RosALES 2005), C. l. concolor chroughour rhe Yucatan Peninsula; both subspecies appear to be most common in coastal areas. Conophis L. dunni is found along rhe Pacific coastal plains ofChiapas (k. VAREZ DEL TORO 1973, 1982, JOHNSON 1989). WELLMAN (1963)
Species Accounts: Alechinophidia: Dipsadidae
considered rhe populations fromChiapas a distinct species, Con ophis pulcher, bur this species is a synonym of C. lineatus according co WILSON & MEYER (1985). The road guarder is a lowland species commonly found ar elevations below 300 m; in central Veracruz, however, this species has been reported from elevations of up co 2,000 m. Natural history: This terrestrial, diurnal snake inhabits tropical deciduous forest and savannas. Ir mostly hums lizards, especially whiprails of the genera Aspidoscelis and Holcosus, and less fre quently rodents; occasionally it feeds on ocher snakes as well as on small birds and their eggs; captive-held specimens will also ear frogs and roads (M ITTLEMAN 1944, CAMPBELL r998a). lncidencal notes on reproduction indicate chat females lay clutches of 4-12 eggs between February and April; neonates have been reported from April to June (KOHLER 2001, SoL6RZANO 2004).
225
and concluded that no subspecies should be recognized in C. vittatus. There are 7 (rarely 8) supralabials, 9-10 infralabials, I loreal, 1 preocular, 2 postoculars, 2+2 temporals, 149-181 vencrals and 55-76 divided subcaudals. Distribution: Conophis vittatus occurs on the Pacific versant from Nayarit and extreme southern Durango southward co the Rio Grijalva Valley of centralChiapas (DUELLMAN 1961, 19656, Al..vA REZ DEL TORO 1973, 1982, S ALDANA DE L A R.ivA & PEREZ RAMOS 1987, RAMfREZ-BAUT ISTA 1994, C AS AS-ANDREU et al. 1996, 2004). le also ranges inland through rhe lower and upper reaches of the Rfo Balsas Depression to eastern Michoacan, Morelos and sourh western Puebla (BENiTEz GALVEZ 1997, AGUILAR et al. 2003, G ARciA-VAZQUEZ et al. 2006). The vertical distribution exrends from near sea level co around 1,200 m (WELLMANN 1963). Natural history: This terrestrial and diurnal species inhabits
Conophis vittatus PETERS, 1860 Striped road guarder; guarda caminos listada, escombrera rayada Conophis vittatus PETERS 1860b: 519. Type locality: unknown. SMITH (1941h) designated first Acapuko, Guerrero as the type localiry; SMITH & TAYLOR (1950b) lacer resrricted it to Laguna Coyuca, Guerrero, Mexico. Conophis sumichrastii sumichrastiiCoPE 1876: 137. Type localiry: "Tehuanrepec and Guadalajara", restricted to Guadalajara, Jalisco, Mexico, by SMI�H & TAYLOR (1945). Conophis sumichrastii viduus CoPE 1876: 137. Type locality: Te huancepec, Oaxaca, Mexico. Conophis viduus SMITH 1939a: 31. Conophis vittatus viduus SMITH 1941h: 120. Identification: A rather scour, striped snake wirh a pointed snout; dorsal color pattern consisting of 3 or 4 black stripes; fosr scale row not pigmented; chin and labial borders whire; usually 7 supra labials; dorsal scales smooch and in 19 rows at midbody. Description: TL up co 84 cm. The coloration in chis species is highly variable. The dorsal ground color is white, gray, bluish gray, cream, pale tan or pinkish. Two different color patterns are known, one of which has three, and rhe ocher four black snipes. Specimens from che Isthmus of Tehuancepec region and the Central Depression of Chiapas usually possess three broad stripes. There is a median black stripe chat initiates on rhe snout and is 3 to 4 scales wide at midbody; and chere is a black lateral stripe rhac initiates on the sides of the snout, passing tbrough cl1e eyes and onto the sides of the body, usually covering 3 scale rows. Specimens from the northern part of the range (Durango to Micboacan, Morelos and Puebla) have four black stripes. Both rhe lateral and rhe dorsolateral stripes are between one half scale and two scales wide. The dorsolateral scripes unite on the nape, forming a broad middorsal srripe char covers rhe top of the head and terminates on rhe snout. In borh color morphs, the labials and all ventral surfaces including the first dorsal scale row are im maculate white or cream. Specimens with three scripes have been suggested to be a different subspecies, C. vittatus viduus. WELL MAN (1963) demonstrated a large area of incergradation (from Michoacin somhward coChiapas) benveen the rwo color patterns
tropical deciduous forest. le feeds mostly on lizards (especially Aspidoscelis and Sceloporus) and snakes. Clutches of 5-10 eggs are laid during the dry season berween February and April (A.LVAREZ DEL TORO 1973, RAMIREZ-BAUTISTA 1994).
Genus Cryophis BOGERT & DUELLMAN, 1963 A single species, C. hallbergi, is placed in this genus. This mode rate-sized, semi-arboreal and nocturnal snake is known only from the cloud-forests of northeastern Oaxaca. Based on similarities in overall habicus, vertebrae and the mode of scale-row reduction (23 rows at midbody a□d19 anterior ro vent), B OGERT & DUELL MAN (1963) suggested chat chis species has affinities with snakes of the genera Leptodeira and Ta.nta!ophis. In a study using mrDNA sequence data, however, MULCAHY (2007) found char Cryophis is more closely related co Hypsi.glena and Pseudoleptodeira than it is co Le_ptodeira and lmantodes. Cryophis hallbergi differs from mem bers of the genus leptodeira in having the enlarged rear maxillary reeth ungrooved and the pupil only slightly elliptical. Additional features characterizing the genus include a slender and slightly compressed body; rail medium in length; head wide, flat-copped and well differenciated from rhe neck; snom long and slighcly rrun cace· in dorsal profile; eyes moderately large; basic colubrid comp lement of enlarged head places; nasal undivided; dorsal scales keeled; subcaudals and anal place divided.
Cryophis hallbergi, BOGERT & DUELLMAN, 1963 Hallberg's cloud forest snake; culebra de bosque mes6philo de Hallberg Cryophis hallbergi BOGERT & DUELLMAN 1963: 4. Type locality: "6 kilometers south ofCamparnemo V ista Hermosa, at an ele vation of 1865 meters, ac the norrhern periphery of the Sierra Juarez, Disrriro de Ixclan, Oaxaca, Mexico". Identification: A slender, moderate-sized snake with a dorsal pattern of reddish blotches alternating wirh dark b rown blotc_hes; all blotches extending to edges of ventrals; dorsal scales keeled, in 23 rows ar midbody and 19 rows anterior co vent.
Species Accounts: Alerhinophidia: Dipsadidae
22.6
Fig.
277:
Diadophis punctntttS dugesii, Presa Maria Atizapan, Valle de Mexico, Escado de Mexico, elevation
Fig.
278:
Dipsas brevifacies, Sian Ka'an, Quintana Roo, elevation
10
m.
2,300
m.
P. HEIMES
A. RAMIREZ VELAZQUEZ
Species Accouncs: Alechinophidia: Dipsadidae
Descrjpcion: The maximum TL is abour 75 cm. The dorsal pac rern consists of about 25 reddish brown, saddle-like blotches that alrernace wich dark brown blotches of roughly the same size. The blotches are narrowly separated by pale yellow margins. There is licde uniformity in the shape of the blotches, but che dark brown blocches rend to become narrower on the flanks of che body; Ir regular dark spots are present in the reddish brown areas along che edges of the ventrals. The dorsal surface of che head is dark brown, che sides of che head are mostly pale yellow with a dark ocular stripe excending from che snouc through che eye and then downward co che angle of the mouch. The supralabials and infra labials have dark brown posterior margins, and faint brown dors are scaccered on che ancerior chinshields. The belly is uniformly cream or pale yellow, che subcaudals are increasingly pigmented with brown coward the tip of che rail. The iris is reddish bronze. The holotype, an adulc male, has 9 supralabials, u infralabials, I loreal, 3 preoculars (lower very small), 1-2 poscoculars, 1+2 and 2+2 temporals, 186 ventrals and 67 divided subcaudals. Distribution: Cryophis hal!bergi frequents incermediace elevations (1,000-2,000 m) iJ1 the Sierra Juarez and Sierra Mixe of norrh eascern Oaxaca (BOGERT & DUELLMAN 1963, CASAS-ANDREU et al. 1996). Natural h.istory: This semi-arboreal and nocturnal snake inhabits relatively cool and humid montru1e cloud forest.
Genus Diadophis BAIRD & GIRARD, 1853 As currently understood, the genus Diadophis contains a single species, D. punctatus (BLANCHARD 1942, GEi-rLBACH 1974). Some of che twelve subspecies listed in COLLINS (1990), however, may represent separate species. Diadophis punctatus ranges over much of North America, including extensive areas in northern and central Mexico. This is a secretive, semifossorial snake rarely seen abroad in daylighr. When rhreacened, this inoffensive snake may coil its tail in a spiral and expose the brightly colored underside ro the potential predator. The genus is characterized by the following combination of features: body slender co moderately robust and cylindrical; tail shore ro medium; head narrow and barely discinct from the neck; eyes medium and with a rou11d pupil; denticion opisroglyphous, with enlarged rear maxillary teeth; a generalized colu.brid comp lement of enlarged head places; dorsal scales smooth and in 15 or 17 rows at midbody; subcaudals and anal place divided.
Diadophis punctatus (LINNAEUS, 1766) Ring-necked snake; culebra de collar, coralillo, vfbora collareja, vfbora de panza enchilada CoLuber punctatus LINNAEUS 1766: 376. Type locality: California, USA. Diadophis punctatus BAIRD & GIRA.RD 1853: Ill. Diadophis amabiLis BArRD & GIRARD 1853= u3. Diadophis regalis BAIRD & G1RARD 1853: II5. Type locality: "So nora'', restricted ro Santa Magdalena, Sonora, Mexico, by SMITH & TAYLOR (19506).
Diadophis punctatus laetus ]AN 18636: 262. Diadophis punctatm dougesii VrLLADA 1875: 226. Type locality: Pocreros de Balbuena, near Mexico [Ciryl, Disrriro Federal [Mexico). Diadophis regalis arizonae BIA.NCH.ARD 1923: 2. Diaduphis amabilis simiLis BLANCHA.RD 192r 4. Type locality: San Diego, California, USA. Diadophis anthon)'i VANDENBURGH & SLEVIN 1923: 1. Type locality: South Todos Santos Island, Baja California, Mexico. Diadophis regaLis dougesii TAYLOR & SMITH 1939: 240. Diadophis amabiLis anthonyi BLANCHARD 1942: 47. Diadophis diigesii BLANCHARD 1942: 51. Diadophis regalis !aetus BL ANCHARD 194:i.: 55. Identification: This small snake is grayish co nearly black above. and red, orange or yellow below; an orange, yellow or cream coloured neck ring is usually present; belly usually spotted wirh black; dorsals smooch and in 15 or 17 rows at midbody. Description: Mose adulcs are between 25 and 60 cm, rhe maxi mum-recorded Tl is 87 cm. The dorsum may be gray, blue-gray, olive or slate black. Most individuals possess a conspicuous light collar across che neck just behind the head. The collar may be orange, yellow or cream, and often is edged with black. In the subspecies anthonyi, rega!is and similis the neck ring is ofre.n obscure wirh poorly defined edges or may be obsolete. The cop of the head is generally a darker shade of che color of the back. The supralabials are partly cream or yellow. The belly is yellow orange ro coral red, brightest posteriorly, and usually spotted with black. The bright ventral color is restricted ro the ventrals in D. p. dugesii, while in the other subspecies it extends onco the first dor sal scale row at lease anteriorly. Hatchlings tend co be darker above with a cream or white rather than an orange collar. There usually are 7-8 supralabials, 7-9 (mode 8) infralabials, r lore.al, 2 preocu lars, 2 posroculars, and 1+1 or 1+2 temporals. There are 15 dorsal scale rows, 182-209 ventrals and 48-68 divided subcaudals in simi Lis; 17 dorsal scale rows, 146-206 ventrals and 49-59 subcaudals in dugesii; and 17 dorsal scale rows, 210-239 ventrals and 62-72 subcaudals in regalis. Males have tubercles on rhe scales above the venr (in females sometimes also present, bur less prominent). Distribution: From southeastern Canada through che easrern and southern United Scares south inco central Mexico. In the western United Scates, the discribucion is spotty except for rhe Pacific coas tal region where D. ptmctatus ranges continuously from Washing ton to northern Baja California. Twelve subspecies were described, four of which are found in Mexico (COLLINS 1990). Diadophis p. regalis occurs in the higher reaches of the Sonoran Desert, the northern pares of the Sierra Madre Occidenral and its associated sky islands, the Sierra Madre Oriental and the more mesic areas of the ChihuahuanDesert. Diadophis p. dugesii ranges in the Sierra Madre Occidental from ease-central Sonora and central Cnihua hua southward, and then enrers the southern portion of rhe Mexi can Plateau south ro the Transverse Volcanic Cordillera (Jalisco, northern Michoacan, Escado de Mexico, Disrriro Federal, wesr cenrral Veracruz). Diadophis p. similis ranges through northwestern Baja California south co San Quintin and also occurs on che Isla San Marcfn in the Pacific. Diadophis p. anthonyi is ende®c to Isla Sur of che Islas Todos Sancos .in che Pacific. The vertical dis tribution ranges from near sea level co about 3,200 rn (URIBE-PENA
22.8
Species Accounrs: Alethinophidia: Oipsadidae
et al. 1999), but this snake commonly occurs below 2,300 m. Dist
ribution information for Mexico is given in BLANCH ARD 1942, SMITH 1943a, SMITH & TAYLOR 1945, DUELLM AN 1961, 19656, McCoY 1964, WEBB 1984, TANNER 1985, PtR£z-H1G AREDA & 5MJTH 1991, GrusMcR 2002, SMITH et a!. 2005, LEMos-EsrIN AL & SMITH 2007a, b and DIXON & LEMOS-EsPIN AL 2010. Natural history: The ring-necked snake occurs in a wide variety of habitats ranging &om woodland to desert grasslands. Moiscure at rbe surfaces or below is required. In northern Baja California it is common only in the mesic coastal areas of the Pacific. This semi-fossorial snake often takes refuge in small cracks in the ground or in burrows of ocher animals, but usually it is encoun tered beneath or inside rotten logs, under rocks. boards or orher surface debris. It ears earthworms, slugs, insects and ocher small invertebrates as well as salamanders and frogs, bur in drier habi tats snakes and lizards (including Armiella) constitute the primary food (YAN DENBURGH 1922, GEHLB ACH 1974, FITCH 1975, GRJs MER 2002). Diadophis punctatus is quite active during cool tem peratures and can be found year-round. Maring generally occurs during the cooler months from February to April (Kr.AUBER 1928, BLANCHARD 1942, GRISMER 2002); one, perhaps two clutches of 1-10 eggs are laid in June or July. Hatching occurs after a little more than rwo months and harchlings measure about 100-150 mm in TL. Maruricy is generally reached in about 13-14 months (FITCH 1960).
Genus Dipsas LAURENTI, 1768 Snakes of the genus Dipsas (approximately 30 species, rwo of chem in Mexico) are distributed throughout the lowlands of tropical Mexico and Central and South America (PETERS & OR.EJAS-M1RANDA 1970). In Mexico, this genus has a fragmenced distribution, with D. gaigeae occurring in western Mexico Ualisco and Coli ma) and D. brevifacies in the Yucatan Peninsula. Dipsas is a highly evolved genus of nocturnal and primarily arboreal snakes allied with rhe genera Tropidodipsas of tropical Mexico and Central America, Sibon of Mesoamerica and northern South America, and Sibynomorphus of central and souchern South America. Members of the genus Dipsas are small to moderate-sized snakes with a rather slender, compressed body and a shorr, blunt head that is very distinct from rhe narrow neck. Owing to their specialized diet of snails and slugs, these snakes have developed modifications of the jaw apparatus char include the inward orientation of the reerh on the upper jaw, as well as modification or loss of certain bones and muscular connections design . ed to extract snails from their shells and swallow slippery prey (PETERS 1960, 1964, S AVAGE 2002). Additional features characterizing the genus include a moderately long, prehensile rail; large, protuberant eyes with vertically elliptical pupils; demition aglyphous, with maxillary teeth similar in size; basic colubrid complemenr of enlarged head plates; mental groove absent; dorsal scales smooch and in 13 to 19 rows at midbody; subcaudals divided; anal place undivided.
Key to the Mexican Species of the Genus Dipsas Dorsal scales in 13 rows ac midbody ................... Dipsas gaigeae Dorsal scales in 15 rows at midbody .............. Dipsas bre11ifacies
Dipsas brevifacies (COPE, 1866) Shorr-faced rhirsr snake; caracolera de cara corra, chupa cara coles, madre de hormigas
Tropidodipsas brevifacies CorE 1866a: 127. Type localicy: Yucatan, Mexico.
Dipsadomorus fasciatus BocouRT 1884: 135. Type locality: Yucatan. Leptognathus bre11ifacies MocQuARD 1908: 891. Leptognathus maxillaris WERNER 1909: 279. Type localicy: Tabasco, Mexico.
Sibon brevifacies SMITH 1943a: 470. Dipsas brevifacies SMlTH & TAYLOR 1945: 50. Dipsas maxi!laris SMJTH & TAYLOR 194f 51. Identification: A small, slender snake with a laterally compressed body and a pattern of black and light rings, 9-15 light rings on body and 5-12 on rail; 15 midbody scale rows; 70-101 subcaudals. Description: TL up co about 60 cm; males rend to grow a little larger than females (KOFRON 1982). The color pattern of body and tail consists of alternating light and dark rings. The light rings are red, orange, pink, salmon or whjce; the dark rings are sbiny black and usually are two co three rimes wider rhan the former. There are 9-c5 light rings on the body and 5-12 on the rail. The amerior portion of the head is black, the succeeding nuchal ring may be incerrupted middorsally, either completely or partially, by a protrusion of black pigment from the top of rhe head. The color of the nuchal ring is usually more imense than rhe light body rings. In harchlings and juveniles the light rings are usually white. The number of scales between nostril and eye is highly variable, the total number of preoculars plus loreals may vary between r and 6; there are 8-11 (usually 9) supralabials, 9-t4 (usually u or 12) infralabials, 3 postoculars, 2+2 or 2+3 temporals, 162-180 vent rals and 70-101 divided subcaudals. The vertebral scale row is equal in size or sligbtly wider than the adjacent rows. Distribution: Dipsas brevifacies is endemic to the Yucatan Penin sula. This species is rather common in Yucatan and Quimana Roo, but apparently is rare in Tabasco and Campeche. The ver tical distribution extends from near sea level to about 350 m (KOFRON 1982, LEE 1996, KOHLER 2003). Natural history: The short-faced thirst snake inhabits rbe tro pical semi-deciduous forest and xeric rhorn forest covering the karsr plains and limestone hills of the Yucatan Peninsula. Ic is crepuscular and nocturnal and appears to be most active on rainy nights. Although this snake has many of the adaptations associ ated with an arboreal lifestyle, including a laterally compressed body, slender neck and wide head with enlarged and protruding eyes, it is commonly encountered on the forest floor or on roads ar night. KOFRON (1982) suggested chat Dipsas brevifacies is ter restrial-scansorial rather than arboreal and that it remains under ground most of the time. The diet consists almost exclusively of snails, and during the prolonged dry season from November to May, numerous snails retreat into rhe damp recesses within the limestone topography. The occupation of chis subterranean karst niche would require much climbing and therefore seems advan tageous for a species of arboreal lineage. Females lay 2-5 (usually 2-3) eggs during the early rainy season. Hatchlings or juveniles
Species Accounts: Alerhinophidia: Dipsadidae collected at the end of rhe rainy season had a TL of approximately 190 mm (KOFRON r982).
Dipsas gaigeae (OLIVER, 1937) Gaige's thirst snake; 2icaclinan Sihynomorphus gaigeae OLIVER 1937= 22. Type locality: Paso del Rio, Couma, Mexico. Dipsas gaigeae SMITH & TAYLOR 1945= 5r. Identification: A smaU snake with a pattern of black and light rings, 7-12 light rings on body and 3-5 on cail; usually 13 mid body scale rows; 53-72 subcaudals. Description: This is a relatively robust and shore-railed member of the genus Dipsas reaching about 60 cm in TL; females are a licde larger than males (KOFRON 1982).The color parrem of body and rail consists of alrernaring light and dark rings. The light rings are cream-colored, pale yellow, pinkish or orange dorsally and whitish-yellow vencrally, the dark rings are shiny black. Mose light dorsal scales have black rips.There are 7-12 lighr rings on rhe body and 3-5 on the rail. The light body rings are 4-16 scales wide, and the dark rings may be wider or narrower rhan rhe light rings. The yellow or orange head is marked with a black mask which covers the anterior cop of rhe head including at leasr rhe fr onral and orbir region. There are 7 supralabials, 7-ro infrala bials, 1 loreal, no (rarely r or 2) preoculars, 2 (rarely 1) poscocuJars, 1 or 2 primary temporals and 2, 3, or 4 secondary temporals, 155-169 vencrals and 53-72 divided subcaudals. The vertebral scale row is equal in size or slightly wider than the adjacent rows. Distribution: Western Mexico. Dipsas gaigeae is known from coastal lowlands and foothills (up to 800 m) from Jalisco to western Guerrero (KOFRON 1982, REYES-VELASCO et al. 2008a, C. GR0NWALD, pers. comm.).
229
cape device (specialized pseudaucotomy). The predator apparently becomes disrracred by the reflex thrashing of the fractured rail segmem wh.iJe the snake sneaks away.The tail is nor regenerated, bur because of irs great length ir allows for several breaks (and escapes) over rhe life of an individual (SAVAGE 2002). Other fea rures that characterize the genus include head small, narrow and barely differenriared from the neck; snout long and pointed; lower jaw countersunk; eyes small and with a round pupil; denririon opiscoglyphous, with enJarged rear maxillary reeth; basic colubrid complemenr of enlarged head plates; dorsal scales smooth and in 15 or 17 rows at midbody; subcaudals and anal plate divided. Key to the Mexican Species of the Genus Enulius Dorsal scales in r5 rows at midbody; 150-163 ventrals; 5 supralabials ....................................................... Enulius ofigostichus Dorsal scales in 17 rows at midbody; 165-216 ventrals; 7 supralabials ...................................................... Enulius flavitorques Enuliusjlavitorques (COPE, 1869) Pacific long-tailed snake; culebra coluda del Pacifico Liophisflavitorques COPE 1869a: 307.Type locality: Rio Magda lena, Colombia. Geophis unicolor frscl-lER 1882a: 227.Type locality: Mexico; res tricted co Chilpancingo, Guerrero, Mexico, by SMITH &TAYLOR (19506). Enufius sumichrasti BocouRT 1883: 538.Type locality: Isthmus of Tehuanrepec, Oaxaca, Mexico. Geagras longicaudatus COPE 18846: 162. Emdiusflavitorques DuNN r938: 415. Enufius zmicolorTAYLOR & SMITH 1939: 247. Identification: A delicate pale brownish snake wirh a long body and rail, narrow head and projecting snout; 7 supralabials; 17 mid body scale rows; 165-216 ventrals.
Natural history: This terrescrial and predominantly nocturnal snake inhabits tropical rhorn scrub and palm forests of coastal Description:TL up co 50 cm.The dorsum is slare gray or grayish areas as well as humid broadleaf forests in inland river valleys brown and the belly is cream, rhis color extending lateraUy onro (KOFRON 1982). The annual activity is mosdy restricted to rhe the second or chi rd dorsal scale row. Specimens of Enulius flavi rainy season from July ro October.The diet consists of snails and torques from Mexico are characterized by rhe absence of a pale soft-bodied insects (HARRIS & SIMMONS 1967, RAMfREZ-BAUTJSTA nuchal collar (usuaUy present in specimens from Central America). r994). Clutches of 2-5 eggs are laid during the rainy season.Their The snout is distinctly convex in the subspecies unicolor, and incubation lasrs about 100 days, and hatchlings measure abour nearly straight in sumichrasti; rhe rosrral is somewhat enlarged and projecting.There are 7 supralabials, 6-8 (usually 7) infralabials, 160 mm iJ1 Tl (HALE 1977, SHEEHY et al. 2012). 1 loreal, no preocuJar, 1 supraocular, 1-2 postoculars, 1+2 tempo rals, 165-216 ventrals and 85-121 divided subcaudals. Genus Enulius CoPE, 1871 Distribution: Low and moderate elevations (up ro 1,800 m) on This genus contains five species of very thin and elongate, semi rhe Pacific versanc from western Mexico southward to Colom fossorial snakes that are collectively distributed from Mexico to bia.Three subspecies are recognized, rwo of which are found in Colombia; two species occur in Mexico (SMITH et aL 1967). Erm Mexico. Enuliusf unicolor ranges west of the Isthmus ofTehuan lius is one of the several snake genera (including Pliocercus and tepec from western Jalisco southward to southern Oaxaca; and Scaphiodontophis) chat possess a disproportionately long (about 30- from rhe Pacific coasr into rhe lower and upper reaches of the Rio 40% of coral length in Emdius) and robust but fragile cail, and Balsas Basin eastward ro southwestern Puebla (SMITH &TAYLOR about half of the specimens of Enulius char are found have a por 1945, DAVIS & DIXON 1959, DUELLMAN 1961, SALDA NA DE LA RlvA tion of the rail missing.The tail loss appears to be a predatory es- & PEREZ RAMOS 1987, CASTRO-FRANCO & BUSTOS ZAGAL 1994,
230
Species Accounts: AJechinophidia: Dipsadidae
Fig. 279: Dipsas gaigeae, Ixlahuacan, Colima. C. RODRIGUEZ
Fig. 280: Dipsas gaigeae, Colima. C. GRUNWALD
& J. JONES
Fig. 281: Dipsas gaigeae, Zihuacanejo, Guerrero. C. GRUNWALD & J. )ONES
Species Accounts: Alcrhinophidia: Dipsadidac
CASAS-ANDREU et al. 1996, GARciA-V AZQUEZ et al. 2006). Enulius
f mmichrasti occurs east of the Isthmus along the Pacific low
lands of extreme southeastern Oaxaca and Chiapas. This sub species is also found on the Atlantic versanr in the Rio Grijalva Depression of central Chiapas (ALVAREZ DEL ToRO 1973, 1982, JOHNSON r989, KOHLER 2003). Natural history: This nocturnal, terrestrial and semi-fos sorial snake inhabits savannas, tropical thorn scrub, deciduous and semi deciduous forest and pine-oak forest. Individuals usually hide in rotten logs, under rocks or in subterranean cavities, especially in termite nests. The diet consists of termites, anrs and reptile eggs. Females lay one or two eggs.
Enulius oligostichus SMITH, ARNDT & SHERBROOKE, 1967 Mexican long-tailed snake; culebra coluda mexicana Emdius oligostichus SMITH, ARNDT & SHERBROOKE r967: 1. Type locality: "Mexico Highway 54 between San Blas, Nayarit, and Mexico Highway 15 (5-15 miles NE San Blas)".
231
Identification: A delicate pale brownish snake with a long body and tail, narrow head and projecting snout; 5 supralabials; 15 mid body scale rows; 150-163 ventrals. Description: TL up to 34 cm. The dorsum of the body is uni form gray brown or tan, becoming lighter on the sides; the pos terior edges of each scale are darker. The head is brown, a little darker than the body. The ventral surfaces are light gray or cream, this color extending onto the second dorsal scale row. The rost ral is moderately enlarged. There are 5 supralabials, 5-6 infralabi als, 1 loreal, no preocular, I supraocular, 1 postocular, 1+1 tempo rals, 150-166 ventrals and 82-88 divided subcaudals. Distribution: Western Mexico. Emdius oiigostichus ranges along the Pacific coast from north of Mazatlan, Sinaloa, through Naya rit south to the vicinity of Puerto Vallarta, Jalisco (SMITH et al. 1967, McDrARMID & BEZY 1971, PONCE-CAMPOS et al. 2003). Toward the east it follows low elevations into the barrancas of Nayarit and Jalisco as far as Zapopan in the city of Guadalajara. The vertical distribution extends from near sea level up to about 1,550 m (C. GRUNWALD, pers. comm.). Natural history: This secretive, semi-fossorial snake inhabits tro pical deciduous forest. It presumably feeds on ants and termites and lays eggs.
Fig. 282: Enulius jlavitorques, female, Agua del Obispo, Guerrero, elevation 1000 m.
P. HEtMES
2.32
Species Accounts: Alerhinophidia: Dipsadidae
Genus Geophis WAGLER, 1830 T his large and diverse genus is composed of at least 49 species of small snakes char are highly secretive and essentially fossorial. Geophis ranges from Tamaulipas and southern Chihuahua south ward chrough Cemral America to Colombia (DOWNS 1967, PETERS & OREJAS-MlRANDA 1970, WILSON & TOWNSEND 2007, PAv6N-VAZQUEZ et a/. 2013); in Mexico, the genus is represented by at least 32 species. Mose species are found in mow1tainous regions at moderate ro intermediate elevarions (500-2,200 m). These inoffensive snakes feed primarily on earthworms, although their diet also includes slugs and soft-bodied arthropod larvae (CAMPBELL et al. 1983, CADLE & GREENE 1993). All Geophis spe cies are oviparous. Members of rhe genus are characterized by rhe following com bination of features: body slender co moderately robust, cylin drical; rail shore to moderate; head narrow, conical, with little if any constriction at the neck; snout medium to long, poimed ro bluntly rounded, projecting well beyond lower jaw; eyes small and with a round pupil; dentition aglyphous, with maxillary variable in size; basic colubrid complement of enlarged head places, bur internasals fused with prefronrals, or supraoculars absent, in some species; a long loreal separating nasals from orbit; no preocuJar; parietal and supralabial(s) in contact or separated by anterior temporal; dorsal scales smooth or weakly keeled, in 15 or 17 rows throughout length of body; subcaudals divided; anal plate un divided.
Key to the Mexican Species of the Genus Geophis ra. Anterior temporal present .................................................. 2 16. Anterior temporal absent .................................................. 7 2a. Dorsal scales in 15 rows throughout body .......................... 3 26. Dorsal scales in 17 rows throughout body .......................... 4 3a. Venrrals immaculate .............................. Geophis maculiferus 36. Ventrals each divided into dark anterior portion and pale posterior portion .................................... Geophis incomptus 4a. Postoculars two ................................................................. 5 46. Postoculars single .............................................................. 6 5a. Dorsum pinkish yellow wirh series of irregular, transverse brown spocs; veneer pale with numerous reddish brown spocs ................................................................. Geophis isthmicus 56. Dorsum reddish brown with narrow pale crossbands; venter immacuJate creamy white ................... Geophi.s omiltemanw 6a. Venter of body white to cream anteriorly, grading posteriorly co grayish brown with pale posterior borders on venuals ...... ............................................................. Geophis latifrontalis 66. Veneer of body black, checkered with red or yellow blotches in juveniles ........................................... Geophis mtttitorques 7a. Dorsal scales in 17 rows throughout body .......................... 8 76. Dorsal scales in 15 rows throughout body ......................... 22
8a. PosrocuJars absenr ............................................................. 9 86. Postoculars one or two ..................................................... 10 9a. Dorsum slate black ................................. Geophis anocu!aris 96. Dorsum red or white with large black dorsal saddles ........... . .. ...... .. ......... ............... .................. ....... .... Geophis duellmani roa. PosmcuJars cwo .......................................... Geophis bicolor 106. Poscoculars single ............................................................ Jl ua. Dorsal scales smooch throughout body or smooth except above vent, where faintly keeled ................................................ 12 ub. Dorsal scales distinctly keeled on portion of body ranging from posterior portion of body ro all of body except nape ........ 17 12a. Veneer predominantly immaculate white, cream or yellow 13 126. Venter checkered or banded ............................................ 16 13a. lncernasal scales usually fused with prefroncal scales ............. ............. .............. ........... .............. ................ Geophis dubius 136. Incernasal scales not fused with prefroncal scales .............. 14 14a. No supraocular scale present, froneal scale does not encer orbit ........... ....... ...................... .................. ... Geophis rhodoga.ster 146. Supraocular scale usually present, usually distinct from postocular, frontal scale does not enter orbit ............................ 15 15a. Ventrals in females 129-140 ....................... Geophis turbidus 156. Ventrals in females more than 150 ............ Geophis cha!Jbeus 16a. Veneer checkered with yellowish orange and black .............. . ............................................................. , Geophis blanchardi 166. Venter pale brown anteriorly and yellowish white posteriorly ........................................................... Geo phis immarnlatus 17a. Dorsal scales keeled excepr on neck .............. Geophis na.salis 176. Dorsal scales keeled on not more tl1an posterior three-foLLrths of body ........................................................................... 18 18a. Ventral scales brownish black anteriorly, yellowish white posteriorly ..................................................... Geophis carinosus 186. Ventral surface mostly or entirely immacuJace pale .......... 19 r9a. Ventrals 124 or fewer ................................... Geo phis juarezi 196. Ventrals 126 or more ...................................................... 20 20a. Pale nuchal collar absenc ................................................. 21 206. Broad pale nuchal collar present ................ Geo phis occabus 21a. Vencrals 132 or fewer; dorsum uniform gray ......................... ................................................................... Geophis rostralis 216. Ventrals 132 or more; dorsum brown, scales of first row with yellow cencers ............................................. Geophis sieboldi 22a. Dorsal scales weakly to distinctly keeled on at least posterior half of body ................................................................ 23 226. Dorsal scales smooch throughout body or smooch except above vent ...................................................................... 24
Species Accounts: Alerhinophidia: Dipsadidae
233
23a. Pale nuchal collar absent, dorsum uniform dark .................. .......... ............................................................. Geophis sallei 236. Broad pale nuchal collar present, or, if absenr, dorsu.m reddish with dark markings ......................... Geophis laticollar·is
Distribution: Souchern Mexico. Geophis anocularis is known only &om the vicinity ofTorontepec, a village on the Atlantic escarp ment of the Sierra Mixe in east-central Oaxaca at about 1,900 m elevation (CAMPBELL et al. 1983).
24a. Supralabials five .................................. Geophis semidoliatus 246. Supralabials six .............................................................. 25
Natural history: This fossorial snake inhabits humid cloud fo
25a. Dorsum patterned ......................................................... 26 256. Dorsu.m patternless ........................................................ 31 16a. Dorsum brown or black, usually with narrow pale crossbands rescricred co anterior portion of body or rhroughout lengrh of body and rail ............................................... Geophis dugesii 266. Dorsum pale wirh dark markings ................................... 27 27a. Venter immaculate white or cream ................................. 28 276. Veneer nor as above ........................................................ 29 28a. Dorsum cream with reddish brown crossbands ................. .. ............................................................... Geophis canceflattts 28b. Dorsu.m red with black crossbands .............. Geophis juliai 29a. Veneer black with scattered pale blotches ............................ ............................................................... Geophis laticinctt,s 296. Veneer white with black spots ........................................ 30 30a. Dorsum grayish brown with narrow dark crossbands .......... ............................................................. Geophis nigrocinctus 306. Dorsum with irregular dark markings on head and neck, remainder of body dark brown or black ........ Geophis tarascae 3ra. Ventrals plus subcaudals 171 or fewer .......... Geophis pyburni 31b. Ventrals plus subcaudals 175 or more .......... Geophis petersii Geophis anocularis DuNN,
1920
Sierra Mixe earthsnake; minadora de Sierra Mixe Geophis anocularis DuNN 1920: 127. Type localiry: Toroncepec, Oaxaca, Mexico. Identification: A small burrowing snake with a narrow, conical head and a long, pointed snout; dorsum slate-black, veneer red dish pink; a long loreal; prefroncals contacting parietals; no supra ocular; no posrocular; no anterior temporal; dorsal scales in 17 rows, smooth throughout length of body or faintly keeled above venr. Description: TL up ro 29 cm in males and 32 cm in females. This snake has a slate-black dorsum and a reddish pink veneer; scale rows one and two are mottled wicl1 these two colors. The gular area and the distal portion of the rail are suffused with dark pigment. There are 5-6 (mode 6) supralabials, 5-6 (mode 6) infralabials, 122-126 vemrals in males and 123-132 in females, and 35-39 divided subcaudals in males and 29-35 in females.
rest. Individuals were taken from beneath boulders chat were deeply embedded in a grassy meadow (CAMPBELL et al. 1983). Earthworms are the only known prey items. CAMPBELL et al. (I. c.) suggested that eggs are laid at the end of the rainy season in November or December and char hatching occurs at the onset of the following rainy season in May or June. Geophis bicolor GUNTHER, 1868 Mexican Plateau earthsnake; minadora de la Mesa Mexicana Geophis bicolor GUNTHER 1868: 416. Type locality: "from che neighbourhood of Mexico City".The type locality was ques tioned by SMITH & TAYLOR (19506) and regarded as erroneous by SMITH & FLORES-VILLELA (1993). Dirosema bicolor BouLENG.ER (in pare) 1894: 298. Rhabdosoma bicolor COPE (in part): 18856: 385. Catostoma bicolor AMARAL 1930a: r91. Identi-fication: A small burrowing snake with a narrow, conical head and a moderately long , pointed s11our; dorsum black, with pale spots on vemrolateral scales-; a long loreal; rwo posroculars; usually no anterior temporal; dorsal scales in 17 rows, smooch_ rhroughour length or faintly keded above vent; 149-168 ventrals. Description: TL up to 40 cm.The dorsal ground color is irides cent bluish black, with light pin spots in the centers of most scales; the scales on the lower 2-4 dorsal rows have greenish yel low spots.The venter is yellow to greenish yellow, immaculate or spotted with black.The cop of the head is black, with yellowish green mottling on the snout. The supralabials are yellow with black upper margins; the infralabiaJs are yellow, the anterior ones are spotted with. black.There are 6 supralabials, 7 infraJabials, 2 (rarely 1) posroculars, t49-159 ventrals in males and 161-168 in females, and 46-51 divided subcaudals in males and 38-46 in fe males.
Distribution: Wesrern Mexico. Geophis bicolor ranges through the western parr of the Transverse Volcanic Cordillera from sourhern Jalisco and Colima through central Michoacan co south-central Esrado de Me:cico (DOWNS t967, DIXON 1968, LARA GONGORA & FLORES-VILLELA 1980, SMITH & FLO.RES-ViLLELA1993, ROOIU GUEZ-CANSECO et al. 2013).The vertical distribution extends from about 1,800 ro 2,600 m. Natural history: Some individuals of ch.is species were taken from beneath freshly cur logs and boards in logging areas of pine and pine-oak forest (D1xoN 1968).
234
Species Accounrs: Alethinophidia: Dipsadidae
Fig. 283: Geo phis bicolor, Tapalpa, Jalisco.
C. G RUNWALD & J. JONES
Fig. 284: Geop his cancellatu.s, Morelos, Chiapas.
C. G RUNWALD & J. JONES
23 5
Species Accounts: Alechinophidia: Oipsadidae
Fig. 285: Geophis carinosus,
Paso del Moral, Uxpanapa, Veracruz. L. CANSECO-MARQUEZ
Fig. 286: Geophis dubitts,
from 8 km NE ot El Punco, Oaxaca, elevation 2,000 m. P. HEIMES
Fig. 287:
Geophis duellmani,
La Esperanza, Sierra Juarez, Oaxaca, elevation 1,620 m. P. HEIMES
Species Accounts: Alecbinophidia: Dipsadidae Geophis blanchardi TAYLOR & SMITH, 1939 Blanchard's eanhsnake; minadora de Blanchard Geophis blanchardi TAYLOR & SMITH 1939: 245. Type locality: "2 miles SW of Aculczingo [= Cumbres de Acultzingo, now Puerto del Aire), Veracruz [Mexico]". Identification: A small burrowing snake with a narrow, conical head and a moderately long, poinred snour; dorsum bluish gray to brownish black, veneer checkered with yellowish orange and black; a long loreal; one postocular; no anterior temporal; dor sals in 17 rows, smooch throughout length. Description: TL up to about 40 cm. The dorsum is bluish gray or dark brown, the lateral scales are distinctly paler along their posterior margins. The head is mostly grayish, frontal and pari etals are dark brown. The gular region is pale yellow, the infra labials are mottled with gray. The vemer is checkered with aher nating yellowish ro orange and black blotches, the latter may form bands across the veneer. There are 6 supralabials, 7 infrala bials, 1 (rarely 2) postoculars, 150-156 ventrals in males and 159166 in females, and 34-40 divided subcaudals in males and 2834 in females. Distribution: Eastern Mexico. Geophis b!anchardi ranges from the Sierra de Zongolica in west-central Veracruz and southeastern Puebla southward into the Sierra Mazaceca of northern Oaxaca. The vertical distribution extends from about 2,240 to 2,510 m (DowNs 1967, BENITEZ GALVEZ 1997,CANSECO-MARQUEZ eta!. 2004, CANSECO-MARQUEZ & AUSTIN 2005). Natural history: This secretive, fossorial snake inhabits humid pine-oak forest and cloud foresc. Individuals have been found beneath rocks and rotten logs (BENITEZ GALVEZ 1997,CANSECO MARQUEZ et al. 2004).
dorsals. The chin region is dark brown, rhe veneer is cream and immaculate. There are 6 supralabials, 6-8 infralabials, l poscocu lar, 149-171 ventrals and 21-25 divided subcaudals. Distribution: Geophis cancel!att/S occurs on the Pacific versanc of northernCentral America; ic is known only from a few scattered localities along che southern escarpment of che Sierra Madre de Chiapas and mountains of adjacenc southwestern Guatemala. The vertical disuiburion ranges from about 500 to 2,000 m (UNDY et al. 1966, D0\''7NS 1967, JOHNSON 1989. KOHLER 2003). Natural history: Geophis cance!Laws inhabits tropical moncane and lower monrane rain forest. Geophis carinosus STUART, 1941 Keeled earthsnake; minadora carinada Geophis carinosus STUART 19416: 3. Type locality: "Finca San Fran cisco, 27 km NE ofNebaj, El Quiche, Guatemala". Identification: A small, blackish snake wirh a narrow, conical head and a long, pointed snout; a large loreal; no anterior tempo ral; dorsal scales in 17 rows, disci.nccly keeled 011 posterior rwo chirds of body and on rail.
Description: TL up ro about 28 cm. The dorsum is slate-black, rhe posterior edges of che scales in lateral rows are slightly paler. The chin region is brownish and the veneer is cream; each ventral scale has a narrow dark anterior margin and its lateral edges morded wich dark pigment. There are 6 supralabials, 6 infralabi als, a single tiny posrocular, 120-123 ventrals in males and n5-134 in females, and 44-49 divided subcaudals in males and 41-44 in females.
Distribution: Geophis carinatt/S ranges on che Aclanric versanr from che Sierra de Los Tuxclas of southern Veracruz southward co the Sierra de Los Cuchumacanes in Guatemala. In Mexico, this Geophis cancellatus SMITH, 1941 species is known only from the Los Tuxdas region and the north ern highlands ofChiapas (DOWNS 1967, Ar.vAREz DEL ToRo 1973, Chiapas earchsnake; minadora de Chiapas JOHNSON r989, PEREZ-HIGAREDA & SMITH 1991, VOGT et al. 1997). The vertical distribution ranges from about 150 ro 2,000 m, bur Geophis cancellatus SMITH 1941i: r. Type locality: Chicharras, Chi chis snake is commonly found at elevations berween 800 aod apas, Mexico. 1,500 m. A single specimen from che SierraNorce de Puebla as signed ro Geophis carinosus by DoWNs (1967) does not belong to Identification: A small burrowing snake wich a narrow, conical this species according ro PE.REz-HIGAREDA & SMITH (1988) and head and a long, poineed snout; dorsal pattern of dark crossbands NIETO-M ONTES DE OCA (2003). separated by narrow light incerspaces; veneer cream; a long loreal; internasals fused wirh prefronrals; no anterior temporal; dorsal Natural history: Geophis carinosus inhabits tropical moncane and scales in 15 rows, smooth throughout or faintly keeled above venr. lower moncane rain forest, cloud forest and pine-oak forest. Description: A relatively large member of the genus Geophis reaching up to 41 cm in TL. The dorsal pacrern consists of broad black crossbands separated by narrow orange-red bands. There a.re 28-32 black bands on the body and 4-6 complete black rail rings. The black bands cover 3-10 scale lengdis middorsally, rhe red bands cover 1-2 scale lengths middorsalJy and broaden late rally. The fast dorsal scale row is cream, the second may be cream, motcled or dark. The head is black wich a light collar occupying che posterior supralabials, temporal, parietals and one row of
Geophis chalybeus (WAGLER, 1830) Veracruz eanhsnake; minadora jarocha Catostoma chalybeum WAGLER 1830: 194. Type locality: "Mexico". Geophis chalybeus P ETERS 1859: 275. Rhabdosoma guttulatumCoPE 18856: 385. Type localiry: Mirador, Veracruz, Mexico.
Species Accounts: Alerhinophidia: Dipsadidae
2 37
E (lapoides) chalibaeus COPE 18856: 386. Atractus chalybaeus COPE 1900: 12.32.. Identification: A small brownish snake with a small, narrow head and 'a relatively short, bluntly rounded snout; a large loreal; one poscocular; no anrerior temporal; dorsal scales in 17 rows, smooth throughout length. Description: TL up to 30.5 cm. The dorsum is brownjsh, with scales in the lateral rows having a cream-colored central spot; the veneer is yellowish-cream and immaculate. There are 6 suprala bials, 7 infralabials, 1 posrocular, 154-155 ventrals (in females) and 38-41 divided subcaudals (in females). Distribution: Eastern Mexico. Geophis chalybeus is known only from Mirador and Xalapa in west-central Veracruz at about 1,150 m elevation (DOWNS 1967, PEREZ-HIGAREDA & SMITl-1 1991).
Fig. 288: Head of Geophis dubius.
L. CANSECO-MARQUl>Z
Geophis duellmani SMITH & HOLLAND, r969 Duellman's earchsnake; minadora coralilla de Duellman
Natural history: The habitat is cloud forest.
Geophis duellmani SMITH & HOLLAND 1969: 47. Type localiry: "2 miles S Vista Hermosa, 1750 m, Oaxaca, Mexico".
Geophis dubius (PETERS, 1861)
Identification: A small burrowing snake with a narrow, conical head and a long, pointed snout; dorsal pattern of 18-21 dark saddles separated by narrow light interspaces; a long loreal; pre frontals in contact with parietals; no supraocular; no posrocuJar; no anterior temporal; dorsal scales in r7 rows, smooth through out length.
Mesa RZANO 2004). Mating tively dry Pacific coast of western Mexico there are only a few occurs during che end of che rainy season. Like other cat-eyed records from scattered localities (DAVIS & Drxo N 1959, HAR D Y & s·nakes, chis species is oviparous, with reported clutches of up co McDtARMlD 1969, SALDANA DE LA RivA & PEREZ RAMos 1987), nine eggs. A captive female from Cosca Rica produced up co three and in the Yucatan Peninsula ir appears co avoid the drier north clutches per year, which hatched after 62-63 days. The hacch western region (LEE 1996). In most areas, chis snake occurs at lings measured 142-156 mm in TL (G OLDER 1984). low and moderate elevations, but it has a considerable vertical rustribucion in some pares of southern Mexico. Ir has been found at elevations up to 2,200 m in the Sierra Madre del Sur in central Leptodeira polysticta GiiNTAER, 1895 Guerrero, and up to 2,500 m in the Mixceca Alea in Oaxaca (CASAS-A ND REU et al. 1996). Small-spotted car-eyed snake; escombrera manchada, falsa nau yaca, box -kobob (Yucacec Maya), kuyun ka□ (Lacand6n Maya) Natural rustory: This nocturnal snake occurs in a variety of ha bitats, including tropical deciduous forest, evergreen seasonal Leptodeira polysticta G0NTHER 1895: 172. Type locafay: Mexico forest, lowland and moncane rain forest, and pine-oak forest. Ir and Cencral America; restricted to Belize, British Honduras is most common in forested habitats, where individuals are often [Belize], by SMITH & TAYLOR (19506). seen climbing in bushes or trees along screams. During the day, Leptodeim albofusca BouLENGER 1896 (in pare): 95. they hide in bromeliads, tree hollows or under loose bark, or on Leptodeira annulata polysticta DuNN 1936: 693. the ground under logs or piles of dead leaves. This car-eyed snake Leptodeira annulata taylori SMITH 194rp: n5. Type locality: Oriis frequently encountered around breerung aggregations of certain zaba, Veracruz, Mexico. ,• ,,, uee frog species of the family Hylidae, its principal prey. Tc is Leptodeira septentrionalis polysticta Duiii�, r958c: 72. known co feed on the eggs of che red-eyed tree frog, Agalychnis ldentification: A slender, laceraUy compressed snake y,rith 38-70 callidryas, which are laid on leaves above water (D UELLMAN 1958c, small blotches char are as wide as or wider t�Jn long; two lateral CAMPBELL 1998a, So L6RZANO 2004). Ocher prey items includ. e nuchal blotches, sometimes fused posreWl�rly; a median nape lizards, especially anoles, red coffee snakes (Ninia sebae) and sala stripe; vertical pupils; usually 3 preoculars; dorsals in 23 (occa manders. Clutches of 6-12 eggs are lrud during the lace dry season (February co May). Egg incubation ranges from 70 co 90 days, sionally 21) rows at mjdbody. and hacchlings measure about 170-210 mm in TL. DUELLMAN Description: This relatively slender and long-tailed Leptodeira (1958c) noted char females are capable of scoring sperm for seve reaches a maximum TL of 106 cm; females tend to grow a licde ral years and still produce fertile eggs. (or 1+1) temporals, 185-196 ventrals in males and females, and 6376 divided subcaudals in males and 54-68 in females.
,,
larger than males. The dorsum is yeUowish brown to grayish tan and has a series of small dark brown blotches that number 38-70 on rhe body and continue as an irregular series of spots on che rail. Ac midbody, the blotches are 1-3 scales long, extending late rally to scale rows 5-7. A series of smaller brown lateral spots is mostly alternating with rhe dorsal blotches. A pair of nuchal blotches, 4-6 scales in length, may fuse with its posterior edges to form a U-shaped mark. The top of rhe head is yellowish brown or can, often with scattered, irregular dark brown spots, especially on che prefroncals, froncal and parietals. A median dark brown streak on the nape begins on the posterior edges of the parietals and extends posteriorly for the length of four or five scales, some times co connect with the fused nuchal blotches. A chin, brown
Leptodeira punctata
(PETERS, 1867)
Western cat-eyed snake; escombrera de! accidence C,-otaphopeltis punctata PETERS 1867 ("1866"]: 93. Type locality: "South Africa" (in error), restricted co Mazatlan, Sinaloa, Me xico, by SMITH & TAYLOR (19506). Leptodeira pacifica C OPE 1869a: 310. Type locality: Mazatlan, Sinaloa, Mexico. Sibon pacificum CoPE 1887: 67. Leptodeira punctata BouLENGER 1896: 91.
Species Accounts: Alechinophidia: Dipsadidae
Identification: A small snake with a dorsal pattern of dark spots arranged in 5 or 6 longitudinal rows; vertical pupils; usually 7 supralabi als; dorsal scales in 19 rows at midbody; ventrals less chan 170. Description: TL up to 62 cm; females tend co grow a little larger than males. The dorswn is yellowish ran co light brown or red dish brown with 5 or 6 rows of dark brown or black spots.The largest spots are on the paravercebral and adjacent scale rows; in some individuals these are fused to form a series of transverse bars across che middorswn.The lowermost row of spots is often reduced to dark pigment on the edges of the scales. There is a black nuchal blotch, four co six scales in length, which often is divided along the rnidline.The cop of the head is yellowish cream or tan and usually has dark spots. A dark posrorbital stripe extends from the posterior edge of che eye co the angle of the jaw. The belly is immaculate cream-colored or pale yellow.There are 7 (or 8) supralabials, 9 (or ro) infralabials, 1 loreal, 2 (rarely 1) preocu lars, 2 poscoculars, 1+2 temporals, 150-156 ventrals in males and 151-167 in females, and 59-69 divided subcaudals in males and 54-62 i.n females. Disuibution: Western Mexico. Leptodeira punctata occurs along the Pacific coastal lowlands and adjacent slopes of the Sierra Madre Occidencal from northern Sinaloa southward co northwestern Jalisco, and thence inland to rhe Lago de Chapala in central Ja lisco. The vertical distribution ranges from near sea !eve.I ro about 1,600 m (DUELLMAN 1958c, HARDY & McD1ARMID 1969). Natural history: This common terrestrial, nocturnal snake in habits tropical scrub forest. Ir is known co feed on small frogs, primarily of the genera Craugastor and Leptodactylw. Clutches of 6-11 eggs are laid during rhe rainy season (June ro September) (DUELLMAN 1958c, HARDY & McDIARMID 1969, GOLDBERG 2.004). Hacchlings measure about 137-149 mm inTL (GOLDBERG 2004). Leptodeira septentrionalis
(KENNICOTT,
1859)
Northern cat-eyed snake; escombrera septencrional, falsa nau yaca Dipsas septentrionalis KENNICOTT [in BA1RD] 1859: 16.Type loca lity: Matamoros,Tamaulipas, Mexico, and Brownsville, Came ron Country, Texas, USA; restricted ro Brownsville by SMlTH & TAYLOR (1950b). Eteirodipsas annulata var. septentrionalis JAN & SoRDELLI 1870: pt. I, fig. 2. Sibon annulatum septentrionale CorE 1875: 38. Leptodeira septentrionalis STEJNEGER 1891: 505Leptodeira septentrionalis septentrionalis DUNN 1936: 697. Identification: A moderate-sized snake with a dorsal parte.rn of 20-35 dark blotches usually extending onto che first scale row; lateral intercalary spots usually absent; nape stripe usually present; vercical pupils; three preoculars; dorsal scales in 23 (sometimes 21) rows at midbody; ventrals 181-208.
273
Description:TL up ro 81 cm in males and 99 cm in females.The dorsum has a pale yellow, light can or reddish tan ground color and dark brown ro black saddle-shaped blotches or transverse bands char munber 20-35 on the body and 8-19 on the tail. The blotches are 5-9 scales in length, three co four times the length of rhe dorsal light interspaces, and usually extend onto rhe first scale row, bur sometimes onto rhe lateral edges of che ventrals or only to scale row rwo or three.The anceriormost blotch is always noticeably longer rhan the others. A series of small dark spots is sometimes present on the first and second scale row.The dorsal surface of the head is grayish brown with an indistinct and highly variable pattern of dark brown or black markings. A median dark streak on the nape extends from the posterior edges of rhe pari etals backwards for the length of rwo to five scales. A dark post orbi ral stripe extends from rhe posterior edge of the eye ro the angle of the jaw.The supralabials are brown or gray, with the an terior edges cream.The belly is cream-colored, with brown flecks sometimes present on the lateral edges of the ventrals.There are 8 supralabials, 10-u infralabials, 1 loreal, 3 precoulars, 2 poscocu lars, 1+2 temporals, 181-199 ventrals in males and 185-208 in fe males, and 66-94 divided subcaudals in males and 60-81 in fe males. Distribution: SouthernTexas and northeastern Mexico. Lepto deira septentrionalis occurs on the Atlantic versanc from the lower Rfo Grande of southernTexas and northern Coahuila southward through rhe coastal plains ofTamaulipas ro rhe Rio Panuco in northern Veracruz, and in the Sierra Madre Oriental from central Nuevo Leon southward co Hidalgo. The vertical disrriburion ranges from near sea level to around 2,000 m (DUELLMAN 1958c, MARTIN 1958, LINER et al. 1977, RAMfREZ-BAUTISTA et al. 2010, DLXON & LEMOS ESPINAL 2010, LEMOS-ESPINAL & DLXON 2013 and FARR et al. 2013). Natural history:This nocturnal, terrestrial and arboreal snake in habits subtropical scrub forest, tropical deciduous forest, evergreen seasonal forest and pine-oak forest. Ir has also been found in cleared areas within cloud forest (MARTIN 1958). Ir feeds mostly on frogs and lizards. DITMARS (1945) reported a clutch of 12 eggs. Leptodeira splendida GUNTHER, 1895 Splendid car-eyed snake; escombrera sapera, escombrera ensil lada Leptodeira splendida Gi.iNTHER 189r 171. Type locality: Izucar de Matamoros, Puebla, Mexico. Leptodeira personata BouLENGER 1896: 93 (in part). Leptodeira rhombifera DUNN 1936: 693 (in pare). Leptodeira bressoniTAYLOR 1939a ("1938''): 321.Type locality : Ha cienda El Sabino, 20 miles south of Uruapan, Michoacan, Mex.ico. Leptodeim ephippiata SMITH &TANNER 1944: 131. Type localicy: Agua Mar{n, 8.3 miles west-northwest of Alamos, Sonora, Me xico. Leptodeira duellmani PETERS 1954: 25.Type locality : Coalcoman, Michoacan, Mexico.
2 74
Species Accounts: Alerhinophidia: Dipsadidae
Fig. 331: Leptodeira ptmctata, 20 km S of Cosala, Sinaloa.
Fig. 332: Leptodeira septentrionalis, La Escondida, Nuevo Le6n.
T.
DEVITT
T. BURKHARDT
275
Species Accounrs: Alethinophidia: Oipsadidae
Fig.333:
leptodeira septentrionalis, Rio Apulco, Xocoyolo, Puebla. L.CANSECO-MARQUEZ
Fig. 334:
leptodeim splendida splendida, Tepozdan, Morelos. P. HEIMES
Fig.335:
leptodeim splendida bressoni, Mascara, Jalisco.
C. GRUNWALD & J. JONES
Species Accounts: Alerhinophidia: Dipst1didac
Jdenti.6cation: A small co moderare-sized snake with 18-38 blotch es on che body; laceral incercalary spots and nape srripe present; ornace head pattern on fronral and parietals; vertical pupils; dor sals in 21 rows ar midbody, smooth throughout length or weakly keeled anterior co venc; ventrals 160-184. Description: Mose adults range from 40 co 70 cm in TL, but ex ceptionally large individuals can re.ach 100 cm or more (HARDY & MCDIARMID 1969); females tend co attain a greater size than males. The dorsum is dull grayish brown to creamy ran with 1838 chocolate brown or black blotches on che body and 9-20 on che rail. Additionally there are smaller spots on the sides of che body. A median dark sneak on the nape extends from the pos terior edges of rhc parietals through a light neck collar and, in some specimens, joins the first body blotch. The cop of the head usually is marked with dark Y-, 8-, or similarly shaped ornament occupying rhe frontal and parietal scales, and there is a broad dark posrorbical stripe extending co che neck where it may reach co the first body blotch or not. The belly is whice or cream ance riorly, grading co pale salmon pink posteriorly. Three subspecies are recognized on the basis of differences in color pattern. lepto deim s. splendida has 18-25 body blotches that are rounded but irregular in shape and may be interconnected, buc usually are discrete. The blotches are 4-7 scales in length and only slighrly longer than the light inrerspaces, extending laterally onco scale row 3 or 4. The sides of the body are marked wirh a series of elongate spors. Leptodeira s. bressoni has 32-38 blotches that are outlined with an indistinct light border. The blorches are 3-4 scales in length, roughly as long as the Light incerspaces, and usu ally extend laterally co scale row 5 or 6. Some blotches can be divided along the midline and alternate with one another co form a zigzag pattern. Two or three rows of laceral spots are usu ally present. Leptodeira s. ephippiata has 22-25 blotches or trans verse bands char are faintly outlined wirh black, which, in cum, is indisrincrly bordered with cream. The blotches are 5-6 scales in length, twice the length of the light incerspaces, and extend laterally onto scale row 1 or 2. Elongate brown spots are presenr on the lateral edges of the ventrals and rhe f'usr and sometimes the second scale row. There are 8 supralabials, 9-u (usually 10) infralabials, r loreal, 2-4 (usually 3) preoculars, 2 poscoculars, 1+2 temporals, 160-177 ventrals in males and 160-184 in females, and 76-87 divided subcaudals in males and 70-81 in females. Distribution: Western Mexico. The splendid car-eyed snake oc curs ar 500-1,750 m elevation along che Pacific foothills from ease-central Sonora and southwestern Chihuahua co rhe coastal sierra of Michoacan, and thence into the R!o Balsas Depression as far as southwestern Puebla. The three subspecies are discribu ted as follows: Leptodeira splendida ephippiata occupies the north ern part of the range south co southern Sinaloa; L. s. bressoni ranges through western Jalisco, Colima and Michoacan, and L. s. splendida occupies rhe southeastern part, ranging in che R!o Balsas Depression from northeastern Guerrero and Morelos co southern Puebla. Disuibucion information for the species is given in DuELL\.CAN 1958a, c, 1961, 19656, HAftoy & McDIARMID 1969, TANNER 1985, SALDANA DE LA RrvA & PEREZ RAMOS 1987, BENfTEz G,{I,vEz 1997, AGUILAR et al. 2003, GARCIA-VAZQUEZ et al 2006, FLORES-VILLELA & HERNANDEZ-GARCIA 2006, ENDER SON & BEzv 2007s, LEMOS-EsP1NAL & SMITH 2007a and VALoEZ LA REs & MuN1z-MARTfNEZ 20Ha.
Natural history: Leptodeim splendida occurs primarily in tropical thorn scrub and deciduous forest, bur was also reported from pine-oak forest in Michoac:in (DUELLMAN 1961). This nocturnal snake is usually encountered near water, foraging both on the ground and in trees. le feeds mostly on roads, frogs and egg masses of arboreal frogs; occasionally arthropods are also eacen. DUELLMAN (1958c) reported a female L. s. bressoni laying 9 eggs during the dry season in February.
Leptodefra u1·ibei (RAM(REz-BAUTISTA & SMITH, 1992) Uribe's car-eyed snake; escombrera de Uribe Pseudoleptodeira uribei RAMfREz-BAUT!STA & SMITH 1992: 83. Type locality: "Estacion de Biologfa "Chamela" (EBCH, less than 5 km from the Pacific coast, at approximately 19° 3o'N, 105° 03'W, cl. 10-584 m)", Jalisco, Mexico. Leptodeira uribei REYES-VELASCO & MULCAHY 2010: 99. Identification: A small ro moderate-sized snake with dark blotch es that often are staggered or fused into zigzags; 35-44 blotches on body and 15-22 on rail, rhe former extending laterally co scale rows 2 or 3; light inrerspaces 0.5-2 scales wide; cop of head dark brown or black, with cream or red area bordering parietals and reaching onco neck; dorsaJ scales in 23 rows at midbody. Description: TL up co about 60 cm. The dorsal pattern consists of dark chocolate brown blotches chat are about 3-5 scales in length and usually have rhe lateral ends rounded. The cream colored incerspaces separating the blotches can be traversed by a narrow dark streak. A few incerspaces may be staggered ac the midline. Small lateral incercalary blotches are presenc on rhe lower 2-3 dorsal scale rows, ending on rhe ventrals; otherwise the ven eer is cream. The head is dark brown, with individual head places edged, and somecimes sported, with light cream or pink. The nape and temporal regions are white, cream or red, chis area involving the sides of the parietals but leaving a shore median dark streak extending posteriorly onto the nape. There are 8 supralabials, 910 infralabiaJs, 1 loreal, 2. preoculars, 2 posroculars, 1+2 temporals, 188-200 vencraJs and 64-92 divided subcaudals. Distribution: Leptodeira uribei occurs on rhe Pacific coastal low lands and foothills (up to 600 m) from Jalisco souch co Oaxaca (RAMfREZ-BAUTISTA & SMITH 1992, RAMfREZ-BAUTISTA 1994, MIJANGOS et al. 2006, SJ RIA-HERNANDEZ et a/. 2006, REYES VELASCO et al. 2009, REYES-V£.LAsco & MULCAHY 2010, M£RTZ et al. 2011). Natural history: Leptodeira ttribei inhabits tropical semi-decidu ous forest, apparently preferring relatively humid areas within irs range. This nocturnal and primarily terresrrial snake is mainly active during the rainy season Oune through September). The diet presumably consists of frogs (REYES-VELASCO & MULCAHY 2010). A femaJe collected in July concained six eggs (STREICHER et al. 201l).
2 77
Species Accounts: Alechinophidia: Dipsadidae
Genus Manolepis CoPE, 1885
WEG & OLNER 1940, SMITH r943a, SMITH &TAYLOR 1945, PETERS 1954, DAV1S & DIXON 1959, DUELLMAN 1961, 19656, JOHNSON 1978, 1989, SALDANA DE LA RrvA & PEREZ RAMos 1987, RAMfREz BAuTISTA 1994 and CASAS-ANDREU et al. 1996.
This genus consists of a single species, Mano!epis pumami, which ranges along the Pacific coast from Nayarit south ro western Chi apas. This small, terrestrial snake displays a pronounced sexual dichromatism chat is unique among Mexican snakes. While both genders have the dark dorsal stripe typical of chis species, chis stripe is uniformly dark in males bur bas a pale ceorer in females (WERLER & SMITH 1952, JOHNSON 1978). Ocher defining charac ters include a slender, cylindrical body with a moderately long rail; head rather elongate and barely wider than rhe neck; snout somewhat pointed and upturned; eyes relatively small and with a round pupil; basic colubrid complement of enlarged head places; dorsal scales smooch, in 19 (rarely 17) rows ar midbody and 15 rows anterior to che vent; subcaudals and anal plate divided.
forest, semi-deciduous forest and pine-oak forest. This diurnal, terrestrial snake appears ro be most active during the wet season from June cbrough October. Ir is a constricror and preys mostly on lizards, especially whiptails (Aspidoscelis) and spiny lizards (Sceloporus) (WERLER & SMITH 1952, PETERS 1954, RAMIREZ BAUTISTA 1994, SANCHEZ DE LA VEGA et al. 2012). Females lay dutches of up to ten eggs (R,\MfREZ-BAUTISTA 1994). A juvenile found in Guerrero during the end of the rainy season measured 190 mm in TL (SALDANA DE LA RIVA & PEREZ RAMOS 1987).
Manolepis putnami (JAN, 1863)
Genus Ninia
Thinscaled snake; culebra de escamas delgadas
This genus is composed of nine species of small, semifossorial and predominancly nocturnal snakes that often make up an im portant part of the leaf-litter herpetofauna on coffee plantations, hence rhe English common name "coffee snakes". A Spanish com mon name for these snakes is dormilona ("sleeper") in allusion ro their habit of sometimes remaining motionless when discovered beneath a rorren log or in some other hiding place. If threatened, they will flatten the body dorsoventrally, hide che head under the body and ljft the rail in a spiral (GREENE 1975). Occasionally, they also go through a rather dramatic, "cobra sryle" display in which they raise the head and the anterior rhird or half of the body, angle the head at almost 90 degrees to the axis of the neck, and spread the posterior ends of the mandibles, clearly exposing the pale nape band (HENDERSON & HoEVERS 1977a). In Ninia sebae, a common species in many coffee fincas in southern Mexi co, this display is reinforced by the snake's bright coloration char warns predators that they may be dealing with a venomous coral snake. Despite their elaborate display, these harmless snakes never make any attempt to bite.The genus ranges from San Luis Pocos{ on the Arlanric versant and from Oaxaca on the Pacific versant southward through Central America co Ecuador and rhe upper Amazon Basin (PETERS & OREJAS-MIRANDA 1970, VILLA et al. 1988, SAVAGE 2002, KOHLER 2003).Two species are found io Me xico (BURGER & \'v'ERLER 1954, SCHMIDT & RAND r957). Members of the genus are characcerized by the following com binuion of features: body slender co moderately robust and cy lindrical; rail moderately short to long; head barely differentiated from the neck; eyes medium and wirb a round pupil; dentirion aglyphous, with maxillary teeth similar in size; basic colubrid complement of enlarged head plates; dorsal scales keeled and strongly srriaced, in 17 or 19 rows throughout length of body; sub caudals divided; anal place usually entire.
Dromicus putnami )AN 1863a: 67.Type localiry: San Blas, Nayarit, Mexico. Tomodon nasutus COPE 1864: 166.Type locality: Colima, Mexico. Manolepis nasutus COPE 1885a: 176. Manolepis pumami BouLl!NGER 1896. Identification: A small, slender, cream-colored snake wi rh a broad dark, solid-colored or light-centered middorsal stripe ex rending from head ro rip of rail; usually no loreal; head scales curiously overlapping; dorsal scales usually in 19 rows ar midbody. Description: TL up to 57 cm in males and 72 cm in females.The dorsum is cream-colored or light gray, with a dark middorsal stripe about four scale rows wide. This stripe e.xtends from the top of the head rluoughout the body and rail. 1n males, the dor sal srripe is brown and narrowly bordered with dark brown; in females, the dark brown or black borders are more pronounced and the center of the stripe is a much lighter creamish brown. Females have a series of dark spots on the sides.These spots be come less evident or disappear on rhe posterior part of the body. A venuolareral dark stripe, sometimes reduced to two parallel rows of short streaks, runs the length of the body. In males, che lateral spots and the venrrolateral stripe are faim or absent. Fe males are also darker ventrally than males.The chjo and anterior ventral region of females is usually black; in males, these areas are only slightly darker than the remaining ventral surface which is a light brown.There usually are 8 supralabials, 9 (or 10) infra labials, r (rarely 2) preoculars, 2 (rarely 3) posroculars, 1+2 tempo rals, 165-173 ventrals in males and 174-182 in females, and 74-82 divided subcaudals in males and 57-71 in females. Distribution: Western Mexico. Manolepis putnami occurs ac low and moderate elevations (up ro 1,900 m) on the Pacific versanr from Nayarir south to the Isthmus ofTehuancepec and the adja cent western slope of the Sierra Madre de Chiapas. In the north, this species follows low elevations inro che barrancas of the Rfo Grande de Santiago and has been found as far as Zapopan near Guadalajara, Jalisco (AHUMADA-CARRJLLO et a!. 2007). Distri bution information is given in GAoow 1905, OLIV ER 1937, HART-
Natural history: Manolepis p11tnami inhabits tropical deciduous
BAIRD
& GIRARD, 1853
Key to the Mexican Species of the Genus Ninia Dorsum black with a pale collar; subcaudals more than 72 ........... .................................................................... Ninia diademata Dorsum red, with or without black markings, head black, a yellow co.liar followed by a black one; subcaudals fewer than 72 ......... •• .. •• •• .. • • ................................................................ Ninia sebae
Species Accounts: Alerhinophidia: Oipsadidae
Fig. 336: Leptodeira spLendida ephippiata, Espinazo del Diablo, Durango.
Fig. 337: Leptodeira uribei, Colima.
C. GRUNWALD &
J.
J. ]ONES
REYES-VELASCO
Species Accounts: Alethinophidia: Dipsadidae
279
Fig. 338: Manolepis putnami, male, Monte Bonito, ANP La Sepulmra, Arriaga, Sierra Madre de Chiapas, elevation 436 m. A. RAMiREz VEu(zQuEz
Fig. 339: Manolepis putnami, female, Agua del Obispo, Guerrero. T. DEVITr
Fig.340: Manolepis putnami, female, Agua del Obispo, Guerrero (same individual shown in Fig. 339). T. DEVITT
280
Species Accoum:s: Alerhinophidia: Dipsadidae
Ninia diademata BAIRD & GIRARD, 1853 Ring-nedced coffee snake; dormilona de coHar, miahuacillo
Ninia diademata BAIRD & GIRARD r8n; 49. Type local.i cy: Ori zaba, Veracruz, Mexico.
Streptophorus bifasciatus DuMERJL, BrnRON & DuMERJL 1854: 520. Type localiry: Mexico.
Streptophorus labiosus BocouRT 1883: 550. Type localicy "Guace
mala'', restricted ro "Yepocapa, depc. Chimaltenango", Guate mala, by BuRGER & WERLER (1954). Ninia diademata diademata AMARAL 1930a: 151. Ninia diademata ploraror SMITH 1942h: 153. Type localicy: Du rango, state of Hidalgo, Mexico. Ninia diademata nietoi BuRGER & WERLER 1954: 657. Type loca lity: San Andres Tuxtla, Veracruz, Mexjco. Ninia diademata labiosa BuRGER & WERLER 1954: 660. Identification: A small blackish snake with a complete or inter rupted pale collar; ventral scales with bold, dark markings; dor sal scales suongly keeled and in 19 rows ar midbody.
duras; labiosa ranges on the Pacific side from the Isthmus of Te huancepec sourhward to southwestern Guatemala.
Natural history: The black coffee snake inhabits evergreen sea sonal forest, tropical lowland and monrane rain forest, cloud fo rest, and marginally pine-oak foresr. This secretive and nocturnal snake is generally found in rotten logs or beneath leaf litter or piles of decaying vegetation. Ir feeds almost exclusively on slugs and snails (TAYLOR 1949, SEIB 1985). Clutches of 2-4 eggs are laid during the rainy season.
Ninia sebae (DuMERIL, BrnRON & DUMERIL, 1854) Red coffee snake; dormilona, chac-ib-can (Yucatec Maya), kuyun kan (Lacandon Maya)
Streptophorus sebae DuMERlL, BnmON & DuMERJL 1854: 515. Type
locality: "Mexique", restricted to the state of Veracruz, Me xico, by SCHMIDT & ANDREWS (1936); and to Veracruz, Vera cruz, Mexico, by SMITH & TAYLOR (19506). Elapoidesfasciatu.s HALLOWELL 1855: 35. Type Jocalicy: Honduras. Streptophorus sebtte coLl.ttris ]AN 18626: 27. Type localicy: Mexico. Ninia co/Laris CoPE 1866a: 127. StreptophorttS sebae var. punctufattts BocoURT 1883: 547. Type lo cality: "Guatemala", restricted ro "vicinicy of Quezaltenango on rhe Pacific slope of southern Guatemala'' by SCHMIDT & RA.No (1957). Streptophus atraties var. tlrozii, coLlaris G0NTHER 188s: 101. Ninia sebae DUNN 1935: n. Ninia sebae sebae SCHMIDT & ANDREWS 1936: 169. Ninia sebae morleyi SCHMIDT & ANDREWS 1936: 169. Type locali cy: Chichen Itza, Yucatan, Mexjco.
Description: TL up co 42 cm. The dorsum is predominantly slate black with a yeJJow, orange or cream collar chat usually begins at che poscerior margin of the parietals and often is interrupted middorsally by black pigment. The collar is continuous with the light coloration of the labial scales and chin. The keels of the dor sal scales are often whitish, and each scale of the firsc three dor sal rows may have a distinct cream or yellow center. The belly is whitish or cream wich dark pigment on the lateral edges of each vencral scale and a series of black midvencral markings, ranging from small round spots ro larger blotches chat are about half as wide as the ventrals. The tail usually has a continuous midvent ral stripe along the subcaudals. During the breeding season, nup tial tubercles are present on the chin and snout in males. There are 6 supralabials, 6 infralabials, a single scale in the loreal-pre ocular region, 2 poscoculars, 1+2 temporals, 123-145 ventrals in males and 129-155 in females, and 83-106 divided subcaudals in males and 73-98 in females.
Identification: A small red or reddish brown snake with or with out dorsal black markings; head black with a yellow nuchal col lar followed by a black one; ventral scales immaculate or with scattered dark flecks; dorsal scales strongly keeled and in 19 rows at mjdbody.
Distribution: Ninia diademata occurs on the Aclanric versanr from San Luis Potosi and central Veracruz southward through the base of the Yucatan Peninsula ro central Honduras, and on the Pacific versanr from southeastern Oaxaca to Guatemala (TAYLOR 1949, ALVAREZ DEL TORO 1973, 1982, JOHNSON 1989, PEREZ HJGAREDA & SMITH 1991, CASAS-ANDREU et al. 1996, LEE 1996, BENITEZ GALVEZ 1997, VOGT et al. 1997, MENDOZA-QutJANO et al 2006, GUTIERREZ-MAYEN & SA1.Az,\RA.RENAS 2006, RAMiREz BAUTJSTA et al. 2010, DIXON & LEMOS-ESPINAL 2010, LEMOS ESPINAL & DIXON 2013). This is a foothill species that principally occurs in the coffee life area of the upper tropical zone, around 500-1,300 m in altirude, but locally ascends mount�ns co as much as 2,438 m (BURGER & WERLER 1954). Four subspecies (diademata, !abiosa, nietoi and pforator) were recognized in the latest revision of chis species by BURGER & WERLER (i954); according to WILSON & MEYER (1985), however, these subspecies are not well defined. Of the three subspecies occurring on the Atlantic versanr, plorator is found in southeastern San Luis Porosi and eascern Hidalgo, diademata in central Veracruz, eastern Puebla and northeastern Oaxaca, and nietoi from southern Veracruz ro northeastern Hon-
Description: Adults average between 25 and 30 cm, the maxi mum TL is about 40 cm. This wide-ranging species exhibits con siderable variation in color and pattern (SCHMIDT & RAND 1957, \'3. T hree new water snakes of che genus Nfltrix from Me xico. Chicago Acad. Sci. Nae. Hise. Misc. 126: 1-9. CONANT, R. 195,. Notes on three Texas reptiles, including an addition co the fauna of the srate. Amer. Mus. Novic. 1726: c-6. CONANT, R.196t. A new water snake from Mexico, with notes on anal places an:cican colubrid snake (Pseudo ficimia). Herperologica 26(3): 336-343. HARDY, L. M.1972. A systematic revision of the genus Psmdoficimia (Ser pentcs: Colubridae). J. Herpetol. 6(1): 53-69. HARDY, L. M. 1973. Pseudoficimia. Cat. Amer. Amph. Rept. 146: 1-2. HARDY, L. M. 1975:1. A sysremacic revision of che colubrid snake genus Ficimia.]. HerpetoL 9(2): 133-168.
Licerarure Ci red
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Dipsas 10, 20, 22, 228, 305, 313, 324 biscutttta 193 biscutata var. latifascia r97 brevifacies 228-229, fig. 278, 98m gaigeae 228, 229, figs. 279-281, 98m gemmistrata latistrttta. 264 graci!!ima 264 maxillaris 228 sanniolus 309 septentliona!is 273 splendida 264 Dirosema hicolor 233 col/are 208 omiltemanum 248 Dromicus c!avatits 216 fitlvivittis 292 Godmanni 301 lachrymans 303 laureata 296 margaritiferus 69 omiltemanus 299 ptttnami 277 taeniatus 300 Dryadophis 114 ciiftoni 114 fosciatus 114 melanolomus 115 melanolornus sfevi11i 115 melanolomus stt,arti n5 melanolorm,s tehuanae 115 mclanolomus veraecrucis n5 Dryinus aeneus n8 lltt1'tltltS IT8 Drymarchon 10, 20, 24, 47, 65 corttis 65 corais cleofae 67 corais erehermus 67 comis melanun1s 67 corais o,·i.zabensis 67 corais rubid1ts 67 corais imicolor 67 me/anurtts 65, 67-68, fig. 18, wm melamm,s erebennus 67, 68 melanums melanums 67, 68, fig. 62 melanurus orizahensis 67, 68 melanunu mbidtts 67, 68, fig. 63 melammts unicolor 67, 68 Drymobius 20, 24, GS aurigr.elus 102 boddaerti 115 brunneus 68 chloroticus 68, figs. 64-66, 21m dendrophis 68 latemlis 107 Lemniscatus 150 margaritiferus 68, 69, fig. 67, 22m margaritifentsfiswlosus 69
Index
margaritiferus margaritiftrus 69 margaritiferus occidentalis 69 pulche1rimus 150 taeniatus 111 D1yophis acuminatus u8 Elaphe 135, 151 baiidi n.2 ch!.orosoma 154 emo,yi 122. emoryi meahllmorum 122 flavirufa l35, 138 flavirufaflavii-ufa 135 jl.avinfa mat11dai 135 flavirufa phaescens 138 flavirufas 135 !aeta laeta 122 obsoleta bairdi L22 pardalinus 135 phaescens 138 rosaliae 50 scierotica 5 r subocularis 51 triaspis 154 Elaphis Deppei 131 pleurostictus 131 Elapidae 20, 322 Elapinae 322 Elapochrus deppei 283 Elapoides chalibaeus 237 dugesi 237 fasciatus 280 rostralis 250 sernidoliatus 250 Siebo!di 251 E!apomorph1u michoacanensis 155 Elaps affinis 329 aliem,s 32.9 1ipi.at11s 329 bernadi 328 diastema 329 distans 332 distans var. michoacanensis 332 elegans 333 ephippifer 336 epistema 329 eu1J1xanthus 324 fitzingeri 340 fu!vius var. diastema 329 falvius var. epistema 329 fidvius var. jitzingeri 340 falvius var. hypostema 329 fit!vius var. sappe,·i 329 macgravii var. !atico!la,is 336 michottcanensis 332
nigrocinctus 338 tene1·e 340 Enicognathus annulatus 388 vittatus 289, 292 Enulius 20, 23, 229 flavitorques 229-231, fig. 282, 99m jlavitorques sumichrasti 229, 231 flavitorques ttnicolor 2.29 oligostichus 2.29, 231, 99m sumichrasti 229 1micolor 229 Epicmtes 37 Epictinae 26 Epictia 20, 21, 26, 332 goudotii 26-27, 329, 333, 336, figs. 4-5, 1m
goudotii bakewelli 26, 27 goudotii phenops 27 Epiglottophis deppei 131 fineaticollis 134 Eridiphas marcosensts 260 slevini 260 slevini marcosensis 260 Erythro!tzmprus grammophrys 296 lateritius 216 mel11120cephalus 217 E,yx braminus 35 Eteirodipsas annulata var. septentrionalis 273 Eudryas boddaerti mexicanus r15 pulcherrimus 150 slevini 115 Eimectes 37 Eu,yphoiis mayae 167 Eutaenia aurata 360 ch,ysocephalus 357 cyrtopsis 360 cyrtopsis var. fidvus 365 cyrtopsis subsp. ocellata 360 dorsalis 381 efegans 361 flavifabris 361 godmanii 368 Hammondii 368 insigniarum 36r macrostemma 361 marciana 369 megaiops 361 ornata 381 phenax 384 praeocularis 369 proxima 376
pu!chrilatus 377 rutiloris 376 scalaris 381 sumichrasti 384 Exel.encophis nelsoni 313 Exiliboa 20, 21, 39 placata 39, fig. 23, 7m Exorhina maculata 63 Ficimia 10, 20, 22, 69, 77, 138, 154, 322, 332
cana 77 desertorum 79 .fi'ontalis 138 hardyi 72, fig. 68, 23m lineata 62 olivacea 69, 72, fig. 69, 24m olivacea streckeri 73 ornata 72 pub !ia 69 , 72-73, fig. 70, 25m p11blia publia 72 publia tayiori 72 quadrangularis 79 ramirezi 69, 73, 23m ruspator 72, 73, 23m streckeri 69, 73-76, figs. 71-72, 26m variegata 72, 76, 23m Galeodon annularis 319 Geag,-as 20, 23, 76 frontalis 138 longicaudatus 22.9 redimitus 76-77, 27m Geophidi11m dt1bium 237 Ceophis 10, 20, 22, 2.08, 2s2, 324, 332 annulatm 319 anocularis 232, 233, 100m aquilonaris 240 bicolor 232, 233, fig. 283, rotm blanchardi 232, 236, ro2n1 cance!latus 233, 236, fig. 284, 103m carinosus 232, 236, fig. 285, room cha91beus 232, 236-237, 104m dubius 232, 237, 288, 336, 338, figs. 286, 288, room
duellmani 232, 237, figs. -i.76, 287, room
dugesii 233, 237-240, w4m dugesii aquilonaris 240, fig. 291 dugesii dugesii 240, figs. 289-290 gertschi 174 immacu!attts 232, 240, 105m incompttu 232, 240, 249, ro6m isthmicus 232, 241, 106m juarezi 232, 241, ro5m Juliai 233, 241, 103m laticinctus 233, 241-244, 103m laticinctus albiventris 241
Index laticoffaris 233, 244, ro7rn !atiji-ontalis 232, 244, figs. 292-293, 102m
latifi"ontalis latifrontalis 244 fatifrontalis semirtnnu!atus 2.44 lippiens 167 longiceps 245 maculiferus 232, 245, 106m mutitorques 232, 245, figs. 294-295, 102m
mutitorques yucatanicus 163 nasalis 232, 245, 337, 339, 107m nigrocinctus 233, 248, 249, J04ffi occabus 2.32, 248, figs. 296-297, 107m omiltemana 2.48 omiltemanus 232, 248-249, fig. 298, ro6rn
petersi 249 petersii '233, 249, fig. 299, I04m pyburni 233, 249, ro7m rhodogaster 232, 249-250, room /"Mtralis 232, 250, 105m 1·ussatus 244 sallaei 244, 250 sallaei russatus 144 sallei 233, 250, fig. 300, ro7m semiannulatus 244 semidoliattts 233, 250-2-51, 333, figs. 301302, 103m
sieboldi 232, 251, fig. 303, 107m sieboldii 151 tarttscae 233, 251, fig. 304, 104m tecpanecus 313 turbidus 232, 251, 100m 1tnicolor 229 zebrina 241 Glaphyrophis lateralis 216 pictt1s 212 Glauconia albifrons 16 boettgeri 27 dissecta 29 dulcis 29 humilis 30 myopica 31 Gyalopion 10, 20, 22, 69, 77, 138, 154, 324 atavus 63 canum 77, fig. 73, 28m canus 77 desertorum 79 quadrangula,·e 77, 79, fig. 74, 29m quadrangu!aris 79 quadrangularis desertorum 79 quadrangularis quad,·,mgularis 79 Gyalopium publium 72 Hapsidophrys mexicanus 98 Hemigenius variabi!is 353
Henicognathus annu!atus 388 Godmanii 301 Herpetodryas aestivus u6 flagelliformis 105 margaritifenis 69 Heterodon 10, 22, 252 kennerlyi 252, figs. 305-306, 108m nasicus 252 nasiczts kennerlyi 252. simtts kennerlyi 151 Himantodes gernmi,tratus 264 gracillimus 264 leucomelas 264 tenuissimm 265 Homalocranion bimacu!atum 174 comnatum 171 deppei q8 planiceps 183 Homa!ocranimn atriceps 170 bocourti 171 boulengeri 184 breve 192 calamarinmn 174 canula 192 canulum 192 fiismm 181 Jani r81 melanocephalum var. bocourti 171 michot1caneme 155 miniatum 178, 184 moestum 182 praeoculum 182 rubrum 184 schistosum 184 wilcoxi 189 Homalopsis quinque-vlttatus 220 Homalosoma episcopum 158 HydrophiiMe' 322
Hydrophis 20, 21, 343 platurus 343-344, fig. 409, 151m Hydrops lubricus 220 Hypsiglena 20, 25, 225, 252, 285, 325 affinis 253 109m catalinae 253-256, 261, fig. 308, uom chlorophaea 2.53, 256, fig. 309, 109m chlorophaea chlorophaea 156 ch!orophaea deserticola 256 chlorophaea tiburonensis 256 discolor 312 dunklei 256 gularis 257 jani 253, 256-257, 109m
Jani dunklei 256, 257 Jani Jani 256, 257, figs. 3w-312 Jani texana 256-257, fig. 313 latifasciata 285 mtchalatus 2 57 ochrorhyncha 253, 257, 109m ochrorhyncha baueri 257 ochrorhyncha gularis 257 ochrorhyncha klauberi 257, fig. 3r4 ochrorhyncha martinemis 257 ochrorhyncha ochrorhyncha 257 .. odn-orhyncha venusta 257, fig. 315 ochrorhynchus 257 ochrorhynchus chlorophaea 256 ochrorhynchits deserticofa 156 ochrorhynchus Janii 156 ochrorhynchtts klauberi 257 ochrorhynchus Lorealus 256 ochrorhynchus texana 256 ochrorhynchus tortttgaensis 257 sLevini 253, 260, figs. 316-318, 110m sp. fig. 307
tanzeri 153, 260, fig. 319, mm texana 256 torquata 253, 256, 260-261, fig. 320, 109m
torquata a/finis 253 torquata baueri 157 torquata catalinae 153 torquata dunklei 256 torquata gularis 257 torquata martinensi, 257 torquata tiburonensis 253 torquata torquata 260 1,11aoc1dttnis 253, 261, uom venusta 257 lmantodes 20, 23, 25, 225, 261, 285 cenchoa 26r, 264, fig. 321, 112m cenchoa leucomelas 264 gemmistrat1is 261, 2.64-265, figs. 322323, I13m
gemmistm.tus gemmistratus 165 gemmistratus gracillimus 264, '265 gemmistrattts latistratus 264, 265 gemmistratus lucidoi-sus 264, 265 germnistratus oliveri 264, 265 fatistrahts 264 lucidorsus 264 sp!endidits 264 splendidus l11cidorsus 264 splendidus oliveri 264 splendidus splendidies 264 tenuissimtts 2.61, 265, fig. 324, u4m lndotyphlops 20, 21, 35 1 332 bramimes 35, 329, figs. 14-15, 5m lschnognathits 345 copii 345 dekayi 349 occipitomaculata 352 occipito-maculatt1s 352, 353 storerioides 353
index
Lachesis aurifera 401 barbouri 413 bicolor 404 brachystoma 419, 423 imdulatus 4I8 Lampropeltis IO, 20., 24, 79, 80, 139, )24 abnorma 80-81, figs. 75-76, 30m alterna 80, 81, figs. 77-78, 31m annulata 84, fig. 79, 30m bfairi 81 boy/ii conjunta 84 californiae 80, 84-85, figs. 80-82, 32m californiae californiae 84 califomiae nigrita 84, 92 californiae nitida 84 calligaster So catafinensis 5, 80, 85, 32m doliata abnorma 80 doliata oligozona So doliata polyzont1 89 doliatus schmidti 89 getula 80 getuftts nigrita 84 getulus spiendida 92 greeri 88 herrerae 93 knoblochi 80, 85-88, figs. 83-86, 113, 33m leonis 88 mexicana So, 88, 3u11 mexicana a!terna 81 mexicana blairi 81 mexicana greeri 88, figs. 87-90 mexicana mexicana 88, figs. 92-94 mexicana thayeri 88, figs. 95-97 microphoiis arcifera 89 multicincta agalma 93 multicincta herrerae 93 nitida 84 po!yzona So, 89, figs. 91, 98-103, 30m polyzona blanchardi 80 polyzona polyzona 89 pyromelana 85 pyromelana knoblochi 85 pyromelana woodini 85 rmhveni 80, 89-92, fig. L04, 31m thayeri 88 triangulum So trianguLmn abnorma So triangulum annulata 84 trianguium arcifera 89 triangulum blanchardi So triangulum campbelli 89 triangulum conanti 89 triangulum dixoni 84 triangulum nelsoni 89 triangulum oligozona 80 -triangulum polyzona 89 triangu.lum schmidti 89
triangttlum sinaloae 89 triangulum smithi 89 webbi 80, 92-93, figs. 106-108, 31m zonata So, 93, 33m zonata agalma 93, figs. 109-ur zonata herrerae 93, figs. 1u, u4 zonata pulchra 93 La.mprosoma anmtlatum 55 episcopwn 158 Laticauda 343 coiubrina 343 Laticaudinae 322 Leptodeira IO, 20, 25, 225, 265, 284, 285, 309 albofusca 269, 272 annulata 265 annulata cussiliris 269 annulata polystt'cta 272 annulata taylori 272 bressoni 273 cussiliris 265, 269 discolor 312 duelimani 273 ephippiata 273 ftenata 265, 268-269, fig. 325, II6m fi-enatafi-enata 268 ftenata malieisi 268 ftenata yucatanensis 268 fi-enatum 268 guilleni 285 latifasciata 285 maculata 265, 268, 269, figs. 326-327, 115m my;-tacina. 269 nigrofasciata 268, 269-272, fig. 328, 117m pacifica 272 personata 269, 273 po!ysticta 265, 268, 272, fig. 329, 1t8m punctata 268, 272-273, figs. 330-331, n9m rhombifera 273 septentrionalis 268, 273, figs. 332-333, n8m septentrionalis polysticta 272 septentrionalis septentrionalis 273 septentrionalis yucatanensis 268 smithi 269 splendida 268, 273-276, 120111 splendida brt:ssoni 276, fig. 335" splendida ephippiata 276, fig. 336 splendida splendida 276, fig. 334 torquata 256, 260 torquata torquata 260 uribei 268, 276, fig. 337, u6m yucatanensis mal/eisi 268 yucatanensis yucatanensis 268 Leptognathm aLbocinctus 317
bernoullii 320 brevifacies 228 cuwlliceps 320 dimidiatus 305 Dumerili 319 leucostomus 320 maxillaris 228 philippii 317 sanniolus 309 mnicinctus 320 sexcutatus 319 subamw!atus 316 Leptomicrurus 322 Leptophis 20, 24, 93 aestivus 116 ahaetulla 96, fig. II5, 34m ahaetu!la praestans 96 diplotropis 96-98, figs. 1r6-H7, 35m diplotropis aeneus 96 diplotropis diplotropis 96 diplotropis forreri 96, 98 lateralis 107 majalis 116 margaritiferus 69 mexicanus 96, 98-99, 36m mexicanus mexicanus 98, fig. u8 mexicanus septentrionalis 98, fig. u9 mexicanus yucatanensis 98 modestus 96, 99, figs. 120-121, 37m occidentalis praestans 96 taeniata u1 Leptotyphlopidae 20, 26 Lepcocyphlopinae 26 Leptotyph!ops 10, 26 bakewelli 26 bressoni 27 dissectu.s 29 dugesii 30 dulcis 29 dulcis dissectus 29 dulcis iversoni 31 dulcis myopicus 31 du!cis rubellum 29 dulcis supraocularis 29 goudotii 26 humilis 30 humilis cahuilae 30 humilis chihuahuaensis 31 humilis dugesii 30 humilis !evitoni 30 humilis lindsayi 30 humilis segregus 31 humilis slevini 27 maximus 30 myopica 31 myopietlS dissectus 19 myopicus myopicus 31 phenops bakewe!li 26 phenops phenops 26 segregus 31
Index Lichttnura 20, 21, 39 myrio!epis 42 orcurti 39 roseofasca 42 trivirgata 39, 42, 8m trivirgata bostici 42 trivirgattt roseofilSca 42, fig. 25 trivirgata sas!owi 42, fig. 26 trivirgata trivirgata 42, fig. 24 Liochiorophis n6 vernalis 117 Liophis decorata 293 elapoides diastema 284 jlavitorques 229 godmttni 301 janii 256 kinke!ini 303 !achrymans 303 !aureatus 296 tricinctus 284 vittatilS 292 Loxocemidae 20 Loxocemus 20, 21, 45 bicolor 451 figs. 29-30, 9m sumichrasti 45 Lycodon lyrophanes 196 Lycognathus lyrophanes 196 Lygophis lachrymans 303 Lytorhynchus decurttlttlS 127 Manolepis 20, 25, 277 nasutus 277 putnami 277, figs. 338-340, 12un Masticophis IO, 20, 25, 99, 102 anthonyi 102, fig. 122, 38m aurigu!us 99, 102-103, fig. r23, 39m aurigu!us barbouri 103 barbouri 99, 103, 39m bi!ineatus 99, 103-105, fig. 124, 40m bilineatus slevini III flagelliformis 105 fozge!!um 99, 105-106, 38m fozgellum cingulztm 105, ro6, figs. 125-126 flagellum fi1Liginosus w6 flage!Lt,m lineatulits 105, 106 flagellum piceus 105, 106 flagellz,m testaceus 105, ro6, figs. 127-128 flageflum 11ariolosus ro7 faliginosus 99, 106-107, figs. 129-130, 38m
lateralis 99, 107, fig. 131, 39m lateralis barbouri ro3 melanolomus 115 mentovarius 99, 107-no, fig. 133, 41m mentovarius mentovarius 107, no mentovaritlS striolatzu ro7, 110, fig. 132
mentovarius vario!osus I07, llo omatus Ill ruthveni IIO schorti 99, no, 42m schorti ruthveni 110, fig. u4 schotti schotti 110 semilineatus 103 s!evini 99, Ill, 40m taeniatus 99, u1-II4, figs. 135-136, 42m taeniatus austra!is 1to taeniatus girardi m taeniatus schotti uo Mastigodryas 20, 25, u4 cliftoni u4-II5, fig. 137, 43m melanowmm II4, 115-116, 44m melanolomus melanolomus ll5 melanolomus slevini u5 me/.anolomus stuarti 115, fig. 140 melano!omus tehuanae 115, fig. 138 melanofomus verd.ecmcis 115, fig. 139 Masrigophis laterafis 107 Megalops maculatzlS 269 Mesopeltis sannioftu 309 Microphis quinquelineatus r63 Micmroides 20, 22, 322, 323, 324 euryxanthus 324-325, 146m euryxanthus austra!is 324, fig. 386 euryxanthus euryxanthus 324, fig. 385 euryxanthtlS neglectt,s 324 Micrurus 20, 22, 154, 283, 284, 322, 323, 324, 325, 338
a/finis mayensis 329 bernadi 322, 325, 328, fig. 387, 147m bogerti 325, 328, 147m browni 325, 328--329, 147m browni browni 329, fig. 388 browni taJ,lori 329, fig. 389 diastema 81, 284, 325, 329-332, 147m diastema a/finis 332 diastema alienus 332 diastema apitttus 332, fig. 390 diastema bernadi 328 diastema diastema 329, 332, fig. 391 diastema distans 332 diastema macdougalti 329, 332 diastema michoacanensis 332 diastema proximans 340 diastema sapperi 332 diastema ssp. fig. 392 distans 325, 332-333, 147m distans distans 329, 332, 333, fig. 393 distans michoacanensis 332, 333, fig. 394 distans oliveri 332, 333, fig. 395 distans zweifeii 332, 333, fig. 396 elegans 322, 325, 333, 148m elegans elegans 333
elegans veraepacis 333, fig. 397 eph ippifer 322, 325, 336, 148m ephippifer ephippifer 284, 336, fig. 398 ephippifer zapotecus 336, figs. 399-400 euryxanthus 324 fitzingeri 340 fitzingerifitzingeri 341 fitzingeri microga!bineus 34J fidvim jitzingeri 341 fidvius macu!atus 341 fidviw microgalbineus 341 fidvhlS tener 341 fidvius tenere 341 laticol!aris 325, 336-337, 148m laticollaris laticollaris fig. 401 laticoflaris maculirostris 336 latifasciatus 325, 337, figs. 402-403, r49m
limbatus 284, 322, 325, 337-338, figs. 404-405, 149m limbttttlS fimbatus 337, 338 limbatus spiwsomus 337-338 nebu!aris 325, 338, fig. 406, 150m nigrocinctus 325, 338-340, 150m nigrocinctus browni 329 nigrocinctzts ovandoensis 338 nigrocinctus wagneri 338 nigrocinctus zimilensis 338, 339 nucha!is 328, 337 nuchalis taylori 328 pachecogili 325, 340, r5om proximans 325, 340, r5om tamaulipensis 341 tener 325, 340--341, 150m tenerfitzingeri 341, fig. 407 tener macufatus 34r tener microgalbineus 341, fig. 408 tener tamaulipensis 341 tener tener 341 Mimometopon sappeti 20, Mixcoatlus 20, 21, 413, 4�8 barbouri 413, fig. 498, 179m browni 413, 416, figs. 499, 501, r79m melanurus 413, 416-418, figs. 500, 502-503, 179m
Morenoa orizabemis 67 Naja 322 Nanicidae 7, 20, 345 Natrix IO, 346 bogerti 348 erythrogaster 348 erythrogttster a/ta 348 erythrogaster bogerti 348 jlavirufas 135 rhombifer 348 rhombifera blanchardi 348 rhombifera rhombifera 348 rhombifera werleri 348
Index
sipedon var. erythrogaster 348 storerioides 353 s1'bcarinata 361 taxispilota rhombifara 348 triaspis 154 valida isabeL!eae 385 valida thamnophisoides 385 Nerodia ro, 20, 24, 346, 353 Couchii 348 erythrogaster 346, 348, fig. 4n, 153m erythrogaste1· alta 348 erythrogaster bogerti 348 erythrogaster transvetsa 348 melanogaster 372 rhombifar 346, 348-349 rhombifer blanchardi 349, figs. 4L2-413 rhombiftr rhombifer 349 rhombifer werleri 349 rhombifera blanchardi 348 rhombifera rhombifort1 348 rhombifera werleri 348 rufipunctatus nigronuchalis 373 sipedon rhombifer 348 valida 385 Ninia 20, 22, 208, 277, 332 collaris 280 diademata 277,280, figs. 341-342, 122m dit1demata diademata 280 diademata labiosa 280 diademata nietoi 280 diademata plomtor 280 sebae 272, 277, 280-281, 337 , 339 , figs. 342-343, 123m
sebae modeyi 280, 281 sebae pzmctulata 281 sebt1e sebae 280, 281 sieboldi 251 Notechis ater 322 Ogmius acutus 59 vrzriam 62 Opheodrys 20, 23, 24, 116, 166 aestivus n6, fig. 141, 45m aestivus majalis 116 mayae 167 vernalis I16-117, 45m vernalis blancha,di 117 Ophibolus altemus 81 californiae 84 splendidus 92 triang11/us var. mexicanus 88 triangulus var. zonatus 93 Ophiophagus hannah 322 Ophis californiae 84 Ophryacus 20, 21, 413, 418 melanurus 413, 416, 418
und11latus 392, 413, 418-419, figs. 504509, 180m
Opisthiodon torquatus 208 Oreophis boulengeri 88 Oxybelis 20, 24, II7 aeneu.s n8-u9, figs. 142-143, 145, 46m aeneus aeneus 118 aeneus auratus 118 auratus n8 fulgidus n9, figs. 144, 146, 47m microphthalmus u8 potosiensis II8 Oxy,·hina maculata 63 maculata anomalt:t 63 varians 62 Oxyrhopus 20, 24, 281, 324 baileyi 281 doliatus aequifascia 281 petolt:t 281-283, fig. 344, 124m petola aeqwfasciatus 281 petola sebae 281 petolarius 281 petolarius aequifosciatus 281 proximu.s 209 sebae 28.1 Oxyuramts scutell.ntus 322 Ptmtherophis 2.0, 24, 25, 119 bairdi 119, 122, figs. 147-148, 48m emoryi 119, 122-123, fig. 149, 49m emo,-yi meahllmomm 123 gttttatus 122 Pelamis 343 platurus 343 Petalognathus nebulattts 308 Philothamnus modestus 99 Phimothyra decurtata 127 grahamiae hexalepis 147 hexalepis 147 Phrynonax 20, 24, 123 poecilonotu.s 123-126, figs. 150-152, 50m
poecilonotus a1gus 123, 126 poecilonotm poecilonotus 123, 126 Phyllophilophis aestivns 116 Phyllorhynchits m, 20, 24, 126 arenicola 127 browni 126-127, figs. 153-154, 51m browni browni 126 browni fortitus 126 broumi klauberi 126 decurtntus 127, figs. 155-158, 52m decurtatus arenicola 127
decurtattts decurtants 127 decurtatus non-isi 127 decurtatus nubifus 127 decurtatu.s perkinsi 127 decu1·tatus pore/Ii 127 Pituophis 10, 20, 24, n9, 127 affinis r30 annectens 130 bai,di 122 catenifer 130-131, 53m catenifer affinis 130, 131, fig. 159 catenifer annectens 130, 131, fig. 160 catenifer bimaris 134 catenifer corona/is 130, 1,1 catenifer deppei 131 catmiferfidiginatus 130, 131 catenifer fuliginosus 130 catenifer insitlanus 134 catenifer insularis 134 cate-nifer rutilis 130 catenifer rutilus 130 catenifer sayi 130, 131, fig. 161 catenifer vertebra/is 134 deppei 130, 131-134 54m deppei brevilineata 134 deppei deppei 131, figs. 162-165 deppei gibsoni 134 deppei Jani 131, fig. 166 deppei Lineaticollis 134 deppei var. pholidostictus 131 elegans 47 em01-yi 122 in,,t!anus 134 lineaticol!is 130, 134, 55m lineaticollis gibsoni 134, fig. 168 fineaticoL!is linetrticollis 134, fig. 167 me!anoleucus affinis 130 melanoleucus annectens 130 melanoleucus bimaris 134 melanofeucus var. catenifer 130 melanoleucus corona/is 130 melanofeucus fuliginatus 130 melanoleucus imzda.nu.s 134 melanoleucus sayi 130 melanoleucus var. vertebraLis 134 mexicanus 130 sayi affinis 130 sayi bellona 130 vertebra/is 130, 134-135, figs. 169-170, 1
53m
vertebra/is bimaris 134, 135 vertebra/is insufanus 134, 135 vertebra/is vertebralis 134, 135 Pityophis elegans 47 haematois 134 intermedius 151 Pleiocercus aequalis 284 Pliocerczts 20, 22, 229, 283, 324 aequalis 284
Index
andrewsi 284 andrewsi pacificus 284 bicolor 284 bicolor hobartsmithi 284 deppei 284 e!apoides 283-284, 328, figs. 345-350, r25m e!apoides aequa!is 284 elapoides celatus 284 elapoides diastema 284 elapoides hobartmzithi 284 elapoides laticolfaris 284 e!apoides occidentalis 284 efapoides sa!vinii 284 elapoides schmidti 284 elapoides wilmarai 284 faticollaris 284 wifmarai 284 Polyodontophis annulatus 388 Porthidium 20, 21,396, 409,419 barbouri 413, 416 d1mni 419-422,ftgs. 511-513, r8rm godmani 409 hespere 419, 422, figs. 514-515, 181m melamtmm 416 nasutum 419, 422, figs. 510,516-517, 181m nummifer mexicanum 397 nummifer nummifer 397 nummifer occiduum 400 olmec 396,400 tzotzilorum 412 yucatanictmz 419,423, figs. 518-519, r8rm Procinura 154 aemuia 155 Psammophis flage!Liformis 105 Pseudelaphe 20,24,25, 135 jlavirufo 135-138, 56111 jlavirufo flavirufa 135, 138, fig. 172 jlavirufa matudai 135, 138 flavirufo pmdalina 135, 138,fig. 172 phaescens 135,138, £.g. 173, 56m Pseudoboa peto!a 281 Pseudoficimia 10, 20,23,138,154 frontalis 138-139, figs. 174-175,57m ftontafis hi!toni 139 hiltoni 139 pu!cherrima 139 pulchra 138 Pseudoleptodeira 20,25,225,284 discolor 312 latifasciata 284, 285, figs. 35r-352, 126m uribei 276 Pseustes poecilonottts 123 poecilonotus argus 123 poeci!onotus poeci/onotus 123
Pyrhonidae 45 Ramphotyph!ops braminm 35 Regina valida 385 Rena 10, 20,21, 26,27,325 boettgeri 27,3111 bressoni 27-29, fig. 6, 2m dissecta 27, 29, 2m dugesii 30 dulcis 27, 29-30, fig. 7, 2m d11/cis rubeltum 29 hmnilis 27, 30,340, figs. 8-10,3m humilis cahuilae 30 humilis d11gesii 30 huml!is humilis 30 maxima 27, 30-31, fig. 11, 2m myopica 27, 3.1, fig. 12, 2m myopica iversoni 31 myopica myopica 31 segrega 27,31, 3m Rhabdophis 345 Rhabdosoma bicolor 233 gurrulatum 236 latifrontale 244 llneatum 204 longiceps 245 mutitorques 245 nasale 245 rostrale 250 semidoliatum 250 zebrinum 24r Rhadinaea 10, :w, 23, 285, 301, 304, 309 aemula 300 binfardi 302. bogertomm 288-289, fig. 353, 127m brevirostris 285 crassa 293 cuneata 288,289, 127m decorata 285, 288,289-292,fig. 354, 128m flavilata 285 farbesi 288, 292, 129m fu!vivittis 288, 292-293, fig. 355, 130m gaigeae 288,293, fig. 356, r29m todmani 285,301 Godmanni 301 hannsteini 302 hernpstedae 302 hesperia 7, 288, 293-296,fig. 357, 131m hespe1·ia baileyi 293 hesperia hesperia 293 hesperia hesperioides 293 kanalchutchan 302 kinkelini 303 Lachrymam 303 lateristriga 285 laureata 285,296-297, fig. 358, 132m macdouga!Li 288,297, 127m
marcellae 288, 297-298, fig. 359, r27m montana 288, 298, fig. 360,129m myersi 288,299, 127m omi!temana 288, 299, fig. 361, 130m pinicola 303 posadasi 304 quinquelineata 288,298, 299-300, fig. 362,129m schistosa 304 taeniata 285,288, 300-301, 336, 133m taeniata aemula 300, fig. 363 taeniata taeniata 300, fig. 364· veraepacis 303 vittata 292, 293, 299,300 l?.hadinella 10,20,22, 23,300 donaji 301, 135m godmani 301-302,134m hannsteini 301 1 302,337, 135m kanalchuchan 3or, 302-303, 136m kinke!ini 301, 303, 136111 lachrymans 301, 303-304, 134m posadasi 301,304, 136m schistosa 301,304, 135m Rhadinophanes 20, 25, 304,309 momico!a 304, 305, 137m Rhegnops sargii 204 visoninus 205 zebrinus 241 Rhinechis elegam 47 Rhinocheilus io, 20, 24,80,139,324 antonii 142 etheridgei 139-142, fig. r76,58m lecontei 139, 142-143, 58m lecontei antonii 139, 142, figs. 177-178 fecontei clams i39, 142 lecontei etheridgei 139 lecontei !econtei 139, 142, fig. 179 lecontei tessellatus 139, 142, fig. r8o tesse!latus 142 Rhinostoma occipita!e 55 SaLvadora 20,z5, r43 bairdii 143-146,147, figs. 181-183, 59m bogerti 143 decurtata 127 deserticola 143, 146, fig. 184, Gorn grahamiae 146-147,59m grahamiae grahamiae 143,146,147 grahamiae hexalepis 147 grahamiae lineata 143, 146, 147,figs. 185186 grahamiae virgu!tea 147 hexa!epis 143, 147-150, figs. t87-188, Gorn
hexalepis celeris r46 hexaLepis deserticoLa 146 hexalepis hexalepis 147 hexalepis klattberi 147
fndex
hexalepis virgultea 143, 147 intennedia 143, 150, fig. 189, 59m buermedia inte1media r50 intermedia richardi 150 lemniscata 143, 150-151, fig. 190, 61m lineata 146 mexicana 143, 15 1, fig. 191, 6om pulcherrima 150 Sanzinia 37 Scaphiodontophis ro, 20, 23, 229, 324 albonuchalis 388 annulatus 388-389, fig. 461-463, 172.m annulatus annulatus 388 annulatus nothus 388 carpicinctus 388 cyclurus 388 nothus 388 Schmidtophis rubriventris 208 Scolecopbidia 26, 37
Scolecophis aemulus 155 fimziceps 182. michoacanensis 155 scytalinu.s 2.09 Scotophis Il9 bairdi 119, 122 emo,yi 122 mutabilis 154 Senticolis 20, 24, 25, 151 triaspis 151-154, 62m triaspis intennedia 154, figs. 192-194 triaspis triaspis 154 Siagonodon dugesii 30 humilis 30 Sibon 10, 20, 22, 228, 313, 332 annulatum septentrionafe 273 amzulatum yucatanen.sis 268 annuLifera 316 biscutatum 193 biscutatum var. !atifasciatum 197 brevifacies 228 dimidiata dimidiata 305 dimidiatus 305-308, fig. 365, 138m fasciata 316 fasciata fasciata 316 fasciata guen·e1·oensis 316 focheri 317 .fischeri fischeri 3 q ftenatum 268 linearis 305, 308, 138m mystacina 169 nebulata nebulata 308 nebulatus 305, 308-309, figs. 366-367, 139m
11igrofasciarum 269 pacificum 272 penonatum 269 philippii 317
sanniola 305, 309, fig. 368, 138m stl.nniolus 309 sartorii 320 sartorii macdougalli 320 sartorii sartorii 320 sibon 308 sp. 308, fig. 369 upsilon 197 zweifeLi 320 Sibynomorphtts 228 gaigeae 229 Sibynophiidae 20, 388
Sibynophis 388 annulatus 388 Sistmrus 20, 21, 426, 495 catenatus 495 catenatus catenatw 495 catenattts edwardsi 495 catenatus edwardsii 495 edwardsii 495 inte1medi11s 443 miliari11s var. edwardsii 495 ravus 462 ravus bnmneus 462 r,wus exiguus 462 ravus futescens 462. ravus ravus 462 tergeminus 495-503, 198m tergeminus edwardsii 495, 503, figs. 6µ652
Sonora 20, 22, 23, 25, 51, 154, 324 aemufa 154, 155, figs. 195-197, 63m aequalis 158 bmzcroftae 159 episcopa 158 episcopa episcopa 159 ep·iscopa tayfori 159 erythrura 155 michoacanensis 155-158, figs. 198-199, 63m
michoacanensis michoacanensis 155 michoacanensis mutabilis 158 miniata linearis 159 miniata miniata 159 mosaueri 159 mutabilis 155, 158, figs. 200-202, 63m occipitalis 55 occipitafis klauberi 55 semiannulata 154, 155, 158-159, figs; 203204, 63m
semianmtlata bfanchardi 159 semiannulata episcopa 159 semiannulata isozona 159 semiannufata Linearis 159 semiamzulata semianm1lata 159 semirmnulata tayfori 159 tayfori 159 Sonorini 154 Spilotes 20, 24, 47, 68, 159 argus 123
corais ,rielanurus 67 deppei 131 erebennus 67 lineatico!fis 134 melanutus 67 poecilonotus 123 puffatus 159, 162, figs. 205-206, 64-ffi pullatus auribundus 162 pullatus- mexicanus 162 variabilis 162 Stenorhina degenhardti quinquelineata 163 degenhardtii 162 degenhardtii apiata 163 degenhardtiifreminvilfei 163 degenhardtii mexicana 162 freminvillii 163 fteminvi!lii lactea 163 lactea 163 mexicana 162 q1miquelineata 163 Stenorrhina 20, 22, r54, 162, 332 degenhardtii 162-163, 333, fig. 207, 65111
degenhardtii mexicana 162, 163 fteminvillei 163-166, figs. 208-209, 66m
fteminvillii 163 Stenostom4 dttlce 29 Goudotti 26 humile 30 myopicum 31 phenops 26 rubelfum 29 tenuicuium 31 Stilosoma 139 extenuatum 80 Storeria 10, 20, 22, 345, 349 copei 345 decayi var. anomala 349 dekayi 349-352, 155ro dekayi anomafa 352 dekayi temporalineata 349, 352 dekayi texana 349, 352, fig. 414 dekayi tropica 352, fig. 415 hidalgoensis 352, fig. 416, 1_56m occipitomacul.ata 352 occipitomaculata hidalgoensis 352 storei-ioides 353, figs. 417-419, 451, 156m tropica 349 tropica anomala 349 tropica temporalineata 349 tropica tropica 349 Streptophorus atratus var. drozii, coLlaris 280 bifasciatus 280 labiosus 280 sebae 280 sebae collaris 280 sebae var. punctulatw 280
570
Symphimus 20, 23, 166 leucostornus 166, fig. 210, 67m mayae 166, 167, 67m Sympholis 20, 22, 167 lippiens 167-169, figs. 211-212, 68m lippiens lippiens 167 lippiens rectilimbus 167 Tachymenis grammoph,ys 2.96 lateritia 216 melanocephala 217 Taeniophis imperialis 216 Tantalophis 20, 25, 225, 309 discolor 304, 309, 312, figs. 370, 374, 140m Ttmtifla 10, 20, 23, 47, 76, 154-, 169, 189, 191, 322, 325, 329, 332 atriceps 170-171, 179, fig. 213, 69m bimaculat-a 174 bocourti 170, 171, figs. 214-215, 70m bocouni bocourt'i 171 bogerti 189 boulengeri 184 bn:vis 192 brevissima 189, 191 briggsi 169, r71, fig. :1.16, 7Im calamarina 76, 169, 174, fig . 2.17, 72111 canula 189, 192 camt!a brevis 192 ca11ula canu!a r92 cascadae 169, 174, fig. 2r8, 72m ceboruca 169, 175, fig. 219, 72m coronadoi 169, 175, 72m cuesta 181 cuniculator 169, 175-178, 73111 deppei 169, 178, fig. 220, 72111 depressa 76 deviatrix 184 eiseni r83 excubitor 191 flavilineata 170, 178, fig. 221, 73m fi1sca 181 gracilis 170, 178-179, fig. 222, 74m hobartsmithi 170, 179, figs. 223-225, 75m impensa 170, 179-181, 73m jflni 169, 181, 188 janni 181 johnsoni 169, 181-182, 73m kirnia 182 lintoni 189, 192 martindelcampoi 174 miniata 184 miniator 184 moesta 170, 182, 76111 moesta cuniculator 175 moesta moesta 182 morgani 184 nelsoni 313
Index
nigriceps 170, 179, 182, 77m nigriceps kirna 182 nigriceps nigriceps 182 oaxacae 170, 183, fig. 226, 73m planiceps 170, 183, fig. 217, 69m _planiceps at riceps r70, 179 planiceps bogerti 189 planiceps yaquia 189 phrenitica 184 robusta 170, 183-184, fig. 228, 76m ru.bra 170, 184, fig. 229, 78m rub1-a rubra 184 schistosa 170, 184-185, fig. 230, 76m schistosa phrenitica 184 sertula 169, 185, 72m shawi 169, 185, 76m slavensi 170, 187, 71m striata 170, 187, 71 m tayme 169, 187-188, 73m triseriata 170, 188, 71m vu!cani 170, 181, 188, 71m wilcoxi qo, 189, fig. 231, 79m wilcoxi ntbricata 189 wilcoxi wilcoxi 189 yaquia 170, 189, fig. 232, 69m yaquia bogerti r89 yaquia yaquia 189 Tanti!lita 20, 23, 47, 189, 191 brevissima 189, 191, fig. 233, 80111 crmuia 191, r92, fig. 234, 8om lintoni 191, 192, 8om lintoni Linto11i 192 lintoni rozellae 192 Telmraspis nigroads-perms 405 undulatus 418 Thalerophis mexicanus mexicanus 98 richmdi praestans 96 Thamnocenchris aurifer 401 Thamnophis 10, 20, 24, 345, 353 ambdottts 369 baronis-midleri 372 bogerti 356, 357, fig. 420, 157m chrysocephalus 356, 357, figs. 421-422, 158m conanti 356, 360, 157m couchii hammondii 368 cyrtopsis 356, 360 -361, 377 , 159m cyrtopsis col!aris 360, 361, fig. 425 cyrtopsis cyclides 376 cyrtopsis cyrtopsis 360, 361, fig. 424 C)'l'topsis oce!latw 361 cyrtopsis postremtts 376 cyrtopsis pu!chrilatt1s 377 dorsalis C)'clides 360 dorsalis dorsalis 360 dorsalis ocellata 360 dorsa!is _postremus 376
eburatus 357 el egans 357, 361, fig. 426, r58m elegans en-ans 365 elegans hammondii 368 elegans hueyi 361 eques 353 , 356, 361-364, 373, 377, 160m eques carmenensis 364 eques cuitzeoensis 364 eques cyrtopsis 360 eques diluvialis 364 eques eques 364, fig. 427 eques insperatus 364 eques megalops 364 eques obscurus 364, fig. 428 eques patzcuaroensis 364 eques postremus 376 eques scotti 364 eq11es virgatenuis 364 on-am 356, 357, 365, figs. 429-430 , 157m exsuL 356, 365, figs. 43r-432, 157m fulvus 356, 365-368, fig. 433, 161m godmrmi 356, 368, figs. 434-435, 157m halophilus 384 hammondii 356, 368-369, fig. 436, 162m lineri 356, 369, fig. 437, 157m macrostemma 364 marciamts 356, 357, 369-372, fig. 438, 163m ma,-ciamts marciani1s 369, 372 marcianus pmeoczt!aris 369, 372 melanogaste1· 353, 356, 372-373, 164m melanogaster canescens 372, 373 melanogaster chihuahuaensis 372, 373 melanogaster linearis 372, 373 melanogaster mel.anogaster 372, 373 mendax 356, 373, 158m nigronuchalis 356, 373-376, fig. 441. 165m
ordinoides erram 365 ordinoides hammondii 368 ordinoides hueyi 361 phenax 384 phena.x halophilus 384 phenax phenax 384 post,-emus 356, 376, fig. 442, 166m pmeocularis 369 proxima 376 proximus 356, 376-377, 167m proximus alpinus 376, 377, fig. 445 proximus diabolicus 376, 377, fig. 443 proximus orarius 376, 377 proximus rutihris 377, fig. 444 pulchrilatus 356, 377-380, figs. 446447, 166m rossmani 356, 380, fig. 448, 166m rozellae 369 rufipunctatus nigronuchalis 373 mfipunctattts unilabialis 384
Index
sauritu.s chalceus 376 sauritus rntiLoris 376 sca!aris 8, 356, 380-381, figs. 449-451, r68m
sca!aris godnutni 368 scalaris scalaris 380 scalaris scaliger 381 scaliger 356, 381, figs. 452-453, 169m sirtalis 356, 381-384, qom sirtalis dorsalis 38t, 384 sirtalis ornata 381 stejnegeri 364 subcarinata subcarinata 364 sumichra,sti 356, 384, figs. 454-456 , 16un
sumichmsti fulvus 365 smnichrasti praeocularis 369 sumichra.sti sumichrasti 384 unilabialis 357, 384-38 5, fig. 457, 165m valida 385 validu.s 353, 385, 171m validus celaeno 385, 386, fig. 458 validus isabelleae 385 validus thamnophisoides 385, 386, fig. 459 validus validus 385, fig. 460 variabilis 353 vicinus 360 T hrasops mexicanus 98 praestans 96 Toluca 59, 138 amphisticha 62 conica 63 frontalis I38 lineata 62 lineata acuta 59 lineata varians 59, 62 megalodon 63 Tomodon lineatum 224 nasutus 277 Tretanorhinus 7, 20, 24, 201, 312 fateralis 312 nigroluteus 312-313, figs. 371-372, 141m nigroluteus lateralis 312 nigrolutezts mertensi 312 Ii·igonocephalw asper 408 bilineatus 393 nummifer 397 Schlegelii 405 undulatus 418 Trirneresurus aurifer 404 barbouri 413, 416 bicolor 404 brachystoma 423 dunni 419 gm·ciai 416 lansbergii lansbergii 423
rnelanurus 416 nigroviridis aurifer 404 undulatus 418 yucatanicus 423 Trimetopo11 hannsteini 302 posadasi 304 veraepacis 303 Trirnorphodon 10, 20, 24, 25, 192 biscuta.tus 192, 193-196, figs. 235-236, 81m
biscutatus biscutatus 193 biscutatus lambda 196 biscutuams lyrophanes 196 biscutatus semirutus 193 biscutrlttts vandenburghi 196 biscutdtus vilkinsoni 200 collaris 197 fasciofata 197 forbesi 197 lambda 193, 196, fig. 237, 81m lambda lambda 196 lambda paucimacufata 197 lambda paucimacufat11s 197 lambda vilkinsoni 200 !atifoscia 197 Lyrophanes 193, 196-197, fig. 238, 81m major 193 paucimac11!atus 193, 197, fig. 239, 81m quadruplex 192 tau 192, 193, 197-2.00, 82m trl!t latifoscia 197, 200, fig. 243 tau tazt 197, 200, figs. 240-242, 400 tau upsilon 197 upsilon 197 upsilon var. collaris 197 vandenburghi r96 vilkinsonii 193, 200, fig. 244, 81m wilkimoni 200 Tropidoclonion 345 copei 345 ·n-opidoclonium annulatum 317 copii 345 storerioides 353 Tropidodipsas 10, 20, 22, 228, 305, 313, 324
annuli/em 3r3-3r6, fig. 373, 142m bernoul!ii 320 brnvifocies 228 cuculliceps 320 fasciata 313, 316-317, 143m fasciata fasciata 316, 317, fig. 376 fasciata guerreroensis 316, 317, fig. 375 fischeri 313, 317, fig. 377, 144m fischeri kidderi 317 ji-eiae 317 guerreroensis 316 malacodryas 316 occidentala 317
57 1
philippii 313, 317-3-19, figs. 378-380, 143m
repfeta 313, 319, 142m sartorii 313, 319--32.0, figs. 381-382, 142m
sartorii anmtfatw 320 sartorii macdougal!i 320 sartorii sartorii 320 sp. fig. 383
subannulatus 316 zweifeli 313, 320, 142m 'fropidonotus celaeno 385 collaris 360 decayi 349 erythrogaster 348 fasciatus var. erythroga,ster 348 godmani 368 melanogaster 372 mesomefanus 372 proximits 376 quadriserialis 38, rhornbifer 348 scaliger 381 sipedon erythrogaster 348 storerioides 353 tephlopleura 385 transversus 348 validus 385 Tjphlina braminus 35
Typhlopidae 20, 26, 34
Typhlops ba,simacu!atzts 34 braminus 35 microstomrts 34 perditus 34 praelongus 34 tenuis 34
Ungaliophiidae 39 Ungaliophis 20, 22, 39, 42 continentalis 42-43, figs. 27-28, 7m panamensis 42
Uropsophus triseriatzts 478 Urotheca 283 elapoides 234 Virginia fasciata 317
Viperidae 8, 20, 392 Viperinae 392 Xenodon 20, 24, 321 angustirostris 32r mexicanus 321
rabdocephalits 321, fig. 384, 145m rabdocephalus mexicanus p.r Zacholus zonattts 93 Zamenis conirostris 58
5 72
constrictor 58 flage!!iformis 105 latera!is 107 latera!is fuliginosus 106 mexicantts 151 pu!cherrimus 150 taeniatus 111
Index
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