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RE- SQUAMATk SERPENTES: COLUBRIDAE SENTICOLS,S . TIUASPIS Catalogue of American Amphibians and Reptiles. Price, R.M. 1991. Senticolis, S. hiaspis. Senticolis Dowling and Fries SmticoIisDowlingandFries,1 987:202.Type-speciesS, enlicoIis~ Cope 1866, by monotypy. Content. A single species, hiaspis, is recognjzed Definition. Smticolisis a medium-sized, elongate, anteriorly and posteriorly tapered, long-tailed colubrine snake ( s mD owling et al., 1983). The head is narrow, but distinct fromthe slender neck. Theeye issmalltomoderate insizeandwitharoundpupil. Thegenus has a primary row of three elongatetemporals, only two ofwhichmay touch the postoculars (Fig. 1). The tail is equal to 2P2696 of SVL in Figure 1. Head of a juvenile Smticolis t. hiaspis. The three elongate adult females and 30-35so f SVL in adult males. Dorsal scales are primary temporals are characteristic of the species. Courtesy of lightly keeled and have indistinct, paired apical pits. The dorsal scale Herndon G. Dowling and the New York Zoological Society. miaodermatoglyphic pattern is echinate. Scale counts range from 27+35+19t o 33+39+23,w ith many combitions of counts between pointed, and subequal inlength(smaller posteriorly). The prefrontal these exuemes. The midlateral rows are involved in all additions and bone has a pointed anterior projection almost as long as high. reductions. Usually dorsal scale rows VII or WI are involved with The hemipenis (Fig. 2) is extremely distinctive. It is 17-19 counts of 29 or more, and rows V or VI in lower counts. Ventral scales subcaudals in length and subcylindrical in shape. The proximal one- are slightly angulate laterally and range from 241-264 in males and f i io f the organ is smooth. At the distal point of the smooth region 256-282 in females, with females tending to have 15.4-20.5s cales is apair of long, slightly recurved parabasal hooks, one on either side more in different populations. The doacal scute is divided. Paired of the sulcus spermticus. Three to four rows of similar, but smaller, subcaudals range from 87-110 in females and 95-126 in males, with spines occur distal to the hooks and diippear ont he distal half of the mean dimorphism varying 13.2-22.2s cutes in favor of males. organ, where they are replaced by 19-24 oblique rows of papillate The color panem of the crown of the head, though often broken calyces, which deaease slightly insize toward the apex, but continue or obscured in adults, is visible in the young. It is distinaive in having to the end of the organ. The calyces are lozenge-shaped and each of an elongate frontoparietal band with a central opening (Fig. 1). the foursides bears 3-5s tiff papillae. The lips of the sulcus areneither The 19-24 maxillary teeth (usually 21-22) are recurved and raised nor ornamented. Map. Distribution of Senticolis hiarpir. Open cirdes mark known localities, solid symbols mark type-localities for the subspecies. The cross- hatched area indicates a region of intergradation. Diagoosls. Senticolis m yb e distinguished from all other Werner (1 896), Oliver (1937), Hartweg and 0liver (1%0), Taylor and member of the Lampropeltiini( s- Dowling and Fries, 1987) by iu Knobloch (1940), Smith (1943), and Taylor (1951). Subspecific unique hemipenial morphology. With the exception of Asii Elaphe relationships were discussed by Barbour and Cole (1906), Smhh radiata, which differ markedly from Senticolic in morphology and (1941), and Stuart (1948). Descriptions of young were provided by biochemistry, all related genera have clawte or bilobed hemipenes Taylor (1940), Smith (19411, and Woodbury and Woodbury (1944). without distinaive papillate calyces or paired hooks. Additionally, The first occurrence of this species in the United States was docu- Senficolish as a proportionately longer hemipenis than most Elaphe, mented by Stone (191 1). Wright and Wright (1957) gave information and all Pituophis, Bogertophis, and A&ona, the latter genera gen- on identification and coloration, and induded some taxonomic erally possessing organs of 1@16 subcaudals in length. Senticolis notes. Dowling (19%) compared dorsal scale redudom and os- differs from superficially similar members of the Colubrini such as teological features of this species with those of Gonyaroma Shaw Gonyasoma, Aqymgm, and most species of Cobberand Spalero- and Campbell (1974) discussed hei dentification and habitat of the sophis in lacking vertebral or paravenebral scale row reductions. species in Arizona. Lawson and Dessauer (1981) provided data that support the separation of hiaspis from Elaphe, b e d o n electro- Descriptions and Illustrations. See species account. phoretic evaluation of proteins. Dowling and Price (1988) discussed the differences from Bogertophismd the proposed taxonomic status Mstribution. See species account. within the Lampropeltiini Wilson and Meyer (1985) desaibed the species in Honduras, provided a range map, and differentiated the Fossil Record None. subspecies. A brief description, including identification, natural history, and care in captivity of S. hiaspis infennedius appeared in Pertinent Literature. The most important papers con- Mehrtens (1987). The taxon appears in many checklists, keys, notes, cerning this genus are Mertens and Dowling (1952), Dowling (1952, and summary accounts, includmgthoseof Cope(1887,1892), Gadow 1%0), and Dowling and Fries (1987). (1905), ~tqnegerandBarbour(1917),VanDenburgh(1922), Blanchad (1929, Amual(1930), Andrews (1937), Smith (1 938,19431, Hameg Etymology. The name Senticolis is from the Latin sentis, a and Oliver (1940), Smith and Taylor (1945), Perkins (1949), Taylor thorn or bramble, and colis, a penis, in reference to the large spines (19491, Stuart (1950, 19631, Hall (19511, Menens (19524, Manin on the hemipenis. The gender is masculine (Dowlig, pen. comm.). (1955, 1958), Alvarez del Toro arid Smith (1956), Duellman (1957), Cagle (l%8), Behler and King (1979), Lee (1980), Parmerlee and Senticolis triaspis (Cope) Neotropical Ratsnake or Green Ratsnake Coluberrriaspic Cope, 1866:128. Type-locality, 'Belize" (formerly British Honduras), restricted to the city of that name by Smith andTaylor(l950). Holotype, USNM 24903, a juvenile male, col- lected by Dr. Parsons, date of collection unknown (not exam- ined by the author). Ehphe ttiarpis: Amaral, 1930:159. Senricolis mma.pisD: owling and Fries, 1987:202. Content. Three subspecies are recognized: h'aqis, i w - medius, and mutabilk Deflnttion and Magnosis. See the generic account. Descriptiom. The original description was provided by Cope(1866). ~chmidatn dhdrews (1936) gavedetails of scutulation, as did Gaige (1936), who also described the hemipenis. Schmidt (1941) gave the scutulation of the type specimen. Menens and Dowling (1952) and Dowling (1952) provided general descriptions. The most comprehensive desaiption, including meristic chans, il- lustrations, andrange maps, is hat of Dowling (1%0), muchof which is summarized in the reclassificationo f the nenus (Dowlinn and Fries. 1987). A recent description is induded in Wilson and ~ G e(1r98 5): Illustrations. Aphotographofthe typespecimenof Senticolis hiaspis iniemdius appeared in Menens and Dowling (1952). A photograph of a live specimen was included in Dowling (1960), as we1 as an illustration of an evened hemipenis, and dorsal and lateral sketches of the head. Wright and Wright (1957) presented several photographs of S. hiaspis intennediur Color photographs of infennedius are in Shaw and Campbell (1974) and Behler and King (1979). Smhh and Brodie (1982) and Stebbins (1985) provided colored illustrations of in&wnedius. Mehrtens (1987)showed a head close-up of intemdius. A photograph of the miuodermatoglyphic pattern is in Price (1981). Detailed hernipenial illustrations appeared in Dowling and Fries (1987). Distribution. Senticolis hiaspis has a brad geographic dis- tri-bution, nn~in~fromsoutheasteArnri zona andsouthernTamaulipos southw&d th;oigh Mexico and much of Central America to costa Rica. The elevational range is from near sea level to over 2200 m. Fossil Record. None. Pertinent Literature. Much of what has been written about this species concerns taxonomy and distribution The primar~ sources of information are Menens and Dowling (1952), Dowling Figure 2 Right hemipenis of S8nricoli.s hiaqoir, UMMZ 118522,15+ (1952, 1%0), and Dowling and Fries (1987). Earlier references, subaudals in length. The specimens was not completelye vertedand including description of scutulation, indude Boettger (1883), Cope would have extended at least two subcaudals further in life. Courtesy (1885,1900), Bocoun(1888), Boulenger (1894,18%), ~ilnthe(rla %), of F. Waite Gibson. Powell (1980), Ballinger and Lynch (1983), and ~tebbi(i1 985). Etymology. The name hiaspis is derived from the Greek aspis, meaning 'shield," and fmi.s, meaning 'three,' in apparent ref- erence to the three elongate primary temporal head scutes. Remarks. The natural history of this species remains poorly known. The majority of specimens have been taken in montane mesophytic forests along the slopes of the Mexicanhighlands and in Central America. In western and southern Mexico, it may be found in more xeric thorn forest. The elongate habitus, small eyes, and angulate ventrals have led to the supposition hat the species is arboreal, but few sightings are available to support this contention, and many specimens have been collected on roads and trails. The principal prey of S. hiaspisappears to be rodents, although it also eats bids. It has recently been bred in captivity (K. Tepedelen, in litt.). The three subspecies of S. hiaspiswere previously assigned by various authors totally or in part to the three species Elaphe hiaspis, E. cblomoma, and E. mutabilis. Smith (1941) suggested that, based on similaritiesi n scutellation,t he name Cohtber mutabilis Cope was based on an albino specimen of E. hiaspis, and placed it in the synonymy of the latter. Stuart (1948) resurreaed mutabilisas a valid subspecies,r anging fromthephen Forest inGuatemala southwardto Costa Rica on the Caribbean side of Central America. The specimen Figure 3. Smlicdis biaspis inkmnedius from the Chhahua designatedastheh olotypeof Piwphisinknnedius by Boettger(l883) Mountains of Arizona. Photograph by Sam Dunton of a living was shown by Menens and Dowling (1952) to belong to the tuon specimen collected by JamesA . Oliver. previously known as Ehpbe chbmsoma Giinther (1894); thus the latter well-known name, which often appears in the literature, was placed in the synonymy of incennediuc. ined by author). Nahix mutabilk: Cope, 1887:71 (part). Scolopbis mutabilis: Bocourt, 1888:680. Cohberpo~W~e:rn er, 1896247. Type-locality, 'Honduras.' 11- Coluber tria.pis Cope, 1866128. See species account. lustrated on pl. 6, additional type data unknown. Ekapbe hiaspis mutubilis: hart, 1948:68. Nahix hiaspis: Cope, 1887:71. Ehpbe hiaspis in&media.. Mertens, 1952b:93. Scolophis mutabilis: Bocoun, 1888:680 (part). Senticolic hiaspis mutabilk. Dowling and Fries, 1987:202. Ekaphe hiaspis hiaspix Stuart 1948:69. Wticolis hiaspis hiaspit. Dowling and Fries, 1987202, DlqnoeLs. This subspecies is characterized by juveniles with Diagnosis. This subspecies is charaaerized by a median fewer than 80 total blotches, and with a median frontoparietal band frontoparietal band and a s d ,r ounded, pale spot at or near the with an elongate centnl opening. The band is usually broken at the frontoparietal suture. The adults are uniform pale brown dordy. middle of the interparietal suture. A blotched pattern is retained into adulthood. Remark. The hdotype was mistakenly given as USNM 6735 in the original desaiption. Literature Cited Pilyopbis intennediusBoeuger, 1883:148. Type-locality, 'Mexico,' restriaedto Haciend ean Edlt Steanb ibieo&. M ichoadn, Mexico by Dowlinn Alvvez del Toro, M. md H.M. Smith. 19%. Notulae herpetologicae (1960). Holotype, Museum (SMF) 34575, a juvg chiapuhe I. Herpetologka 123-17. nile rmle from the colledion of Dr. Pa~enSte~h.ed ra.t e of col- Amual,A . do. 1930 (1929). Estudos dbre ophidios neotropicos. lection unknown (not examined by ahor). XWI. Lista remissiva dos ophidios da regiiio neou6pia. Mem. Cohtber mubilis: Cope, 1885:175 (part). Inst. Butantan 4:i-vG 129-271. Nahix mutabilit. Cope, 1887:71 (part). Andrews, E.W. 1937. Notes on snakes from theYucatan Peninsula. Scobpbis mutabilis: Bocoun, 1888:680 (put). Cohber cblomsoma Giinther, 1894:115. Type-localities, 'Mexico, Atoyac in Vera Cruz, Amula in Gumero, San Runon" (ldisco), restrictedto Amula (=Almolonga), Gumero by Dowling (1 960). Leaoholotype, British Museum (Natural History) 1946.1.12.99, a w e , collected by H.H. Smirh, date of colleaion unknown (not examined by author). Dowling (1960) cites thir individual as specimen 'b" in Boulenger (1894). It is one of the three in- ferred Giinther cotypes. Ehphe c b b m mS tone, 1911:231. Ehphemutabilis:Taylor, 1%@459.F itu seofthis combination(non C. mutabilis Cope, 1885). Ehphe hiaspis intemedia.. Menens and Dowling, 1952:197. Senticolis hiaspis intermedia: Dowling and Fries, 1987202. Diagnosis. This subspecies is characterizedb y juveniles with a median frontoparietal band with a centnl opening extending forward to the posterior end of the frontal scute, and with 80 or more tod blotches. The adults are unicolor green dordy. 3. Sentko& trfuspfsm utabUls (Cope) Cohrber hiaspis: Cope, 1879:271 (non C. tria.ph Cope, 1866). I I Cohtber mutabilisCope, 1885175 (part). Type-locality, 'Vera Paz,' Figure 4. Subspecific variation in the head pauern of Senticolis Guatemala, restricted to Alta Veravaz hinNands of Guatemala hiaspis as seen in juvenile snakes: a. Senticolis t. triqis, b. S. t. by hart (1948). Holotype, USNM 6745;an adult female, col- mutabilis, c. S. I. inkmnedius Comesy of Herndon G. Dowling and lected by Henry Hague, date of colleaion unknown (not exam- the New York Zoological Society. Zod. Ser. Fld Mus. Nat. Hist. 2&355-359. Co., New York. Ballinger, RE. and J.D. Lynch. 1983. How to know the mphibii Mertens, R 1952a. Die Amphibien und Reptilien von El Salvador, auf and reptiles. Wm C. Brown Co., Dubuque, Iowa. Gmnd der Reisen von R Mertens und A. Zilch. Abh. Sendten- Barbour,T. and LJ. Cole. 1906.Vertebrataf romtheYuatan. Reptilia, bergixhen Naturf. Ges. 487:l-120. Amphibia, and Pisces. Bull. Mus. Comp. Zool. W146-159. -1 952b. Neues iiber die Reptilienfauna von El Salvador. 2001. n Behler,J .L and F.W. King. 1979. The Audubon Society field guide to Anz. 148:87-93. NorthAmerican reptiles and amphibians. Alfred A. Knopf, New -a nd H.G. Dowling. 1952. The identity of the snake AwbLs in- York. tennedius Boettger 1883. Senckenbergiana 33197-201. Blanchard, F.N. 1925. A key to the snakes of the United States, Cana- Oliver,J .A. 1937. Notes on a colledion of amphibii and reptiles da, arid Lower California. Pap. Michigan Acad. Sci., Arts, Let. 4, from the state of Colirna, Mexico. Occ. Pap. Mus. Zd.U niv. pt. Zxiii + 65 p. Michigan 360:l-28. Bocoun, M.-F. 1888. Mission xientifie au Mexique et dans l'Am&- Pumerlee, J.S., Jr. and R. Powell. 1980. Geographicdistribution: Eh- ique Centrale. kpartie. fitudess ur les reptiles a les batrachians. pbe hiaqbir. Herpetol. Rev. 11:116. Livr. 11:657-696. Perkins, C.B. 1949. A key tothesnakesofthe United States. Bull. h l . Boenger, 0.1883. Herpetologixhe Minheilungen. I. Kurze NotiZen Soc San Diego 24:l-79. iiber ReptilienundA mphibien in der HeidelbergerU niversitiits- Price, R.M. 1981.A nalysis ofthe ecologicala ndtaxonomic comspon- Slmmlung. Ber. Offenbacher Ver. Naturk.22:147-152. dence of dorsal snake sale rnicrodermatoglyphics. PhD. the- Boulenger, G.A. 1894. Catalogue of snakes in the British Museum sis, New York Univ. (Natunl History). Vol. 2. London. Schmidt, K.P. 1941. The amphibii and reptiles of British Honduras. -1 896. Catalogue of snakes in the British Museum (Natural Hii- Zool. Ser. Field Mus. Nat. Hist. 20:475-510. tory). Vd. 3. London. -and E.W. Andrews. 1936. Notes on snakes from Yucatan. Zooi. Cagle, F.R 1968. Reptiles, p. 213-268. hr W.F. Blair, A.P. Blair, P. Brod- Ser. Field Mus. Nat. Hist. 20:167-187. korb, F.R Cagle, and GA. Moore, Vertebrates of the United Shaw, C.E. and S. Campbell. 1974. Snakes of the American West. U- States. 2nd ed. McGraw-Hill Book Co., New York. fred A. Knopf, New York. Cope, E.D. 1866. Fourth contribution to the herpetology of tropical Smith, H.M. 1938. Additions to the herpetofauna of Mexico. Copeia America. Proc. Aad. Nat. Sci. Philadelphia 2:123-132. 1938:149-150. -. 1879. Eleventh contribution to the herpetology of tropical -1 941. Notes on Mexican snakes of the genus Ekrpbe. Copeia -1A 8m8e5r.iTc aw. ePlrfothc. c Aonmtreirb. uPthioilnot so. tShoec h. e1r8p:2e6to1l-o2g77y. o f tropical Amer- -11 994413:1. 3S2u-1m36m. vy of the collections of snakes and aocodihns ica. Proc. Amer. Philos. Soc. 22167-1%. made in Mexico under the Walter Rathbone Bacon travelling -. 1887.Catalogueoftheb atrachians and reptiles of Central Amer- scholarship. Proc. U.S. Natl. Mus. 93:393-504. -.i c1a8 9a2n.d A M aietxicicaol .v Bieuwll. o Uf .tSh. eN cahtal.r Maduesr.i s3t2ic:ls- 9a8n.d variations of the -and of NEo.rDth. BArromdirei,a J.r .W 19e8st2e. rAn gPuuibdle. tCoo f.i,e Nlde iwd Ydo~rka. tionr,e ptiles snakes of North America. Proc. U.S. Natl. Mus. 14:58%94. -and E.H. Taylor. 1945. An annotated checklist and key to the -1 900. The a o c d i ,li zards and snakes of North America. snakes of Mexico. Bull. U.S. Natl. Mus. (187)l-239. Rep. U.S. Natl. Mus. for 1898:153-1294. -a nd-. 1950. Typel ocalities of Mexican reptiles andunphibi- Dowling, H.G. 1952. A taxonomic study ofthe American representa- an^. Univ. Kansas Sci. Bull. 33313-390. tives of the genus EhpbeFitzinger, with particular attention to Stebbi,R C. 1985. A fieldguide to western reptiles andmphibii. the forms occurring in Mexico and Central America. Ph.D. the- 2nd ed. Houghton Mifflin Co., Boston. sis, Univ. Michigan. Stejneger,L andT. Barbour. 1917. A checklist of NohAmericanm- n -1 958. A taxonomic study of the ratsnakes. VI. Validation of the - phibii and reptiles. Harvvd Univ. Pr., Cambridge. genera ConysomuWaglera nd ElqbePiinger. Copeia 1958:29- Stone, W. 1911. Onsome collectionso f reptiles arid bamchians from 40. the western United States. Proc. cad Nat. Sci. Philadelphia44: -.1 960.A tuonomicstudyoftheraf~nakesg, enus ElapbeFitzinger. 222-232. VII. The ~issebionh.l ogica 45:53-80. Stuart, L.C. 1948. 'Ihemphibii andreptiles ofAltaVmpaz, Guate- -a nd I. Fries. 1987. A taxonomic study ofthe ratsnakes. VIII. A -m ala. Misc. Publ. Mus. Zool. Univ. Michigan 69:l-109. proposed new genus for Ehpbe tr&ir(Cope). Herpetologia 19%. A geographic study of the herpetofauna of Alta Verapaz, -,4 R3 2H00ig-h2t0o7n. , G.C. Maha, and LR Mwon. 1983. Biochemical -1G 9u6a3te.mA aclhae. cCkolinstt roifb t. hLea hbe. rVpeertto. fBaiuonl.a U ofn Givu. Mateicmhaiglaa.n M4i5s:cl.- P7u7b. l. evaluation of colubrid snake phylogeny. J. 2.001. 201:309-329. Mus. 2001. Univ. Michigan (122):l-150. -and R.M. Price. 1988.A proposed new genus for EIapbedmcu- Taylor, E.H. 1940. SomeMexicanserpents. Univ. Kvlsv Sci. Bull. 26: lank and Ekapbe maliae. The Snake 20:52-63. 445487. Duellman, W.E. 1957. Notes on snakes fromthe Mexican state of Sina- -1 949.A preliminary account ofthe herpetology of the state of la. Herpaologia 13237-240. Sul Luis Potmi, Mexico. Univ. b a sSc i. Bull. 33169-215. Gadow, H. 1905. The distribution of Mexican amphibii and rep- -1 951. A brief review of the snakes of Costa Ria. Univ. Kansas tiles. Proc. 2,001. Soc. London 2:191-244. Sci. Bull. 343-188. Gaige, H.T. 1936. Some reptiles and amphibii from Yucatan and -andl.W. Knobloch. 1940. Repon onanherpetological collection Cunpeche, Mexico. Publ. Camegie Inst. Washington457:289- from the Sierra Madre Mountains of Chh~ahuaP.r oc. Biol. Soc. 304. Washington 55125-130. Giinther, A.C. 1894. Biologia Cedi-Americana. Reptilk and Batra- Van Denburgh, J. 1922. The reptiles of western North America. Occ. chia. RH. Porter, London. Pap. California Aad. Sci. 10:l-1028. Hall, C.W. 1951. Notes on a small herpetological collection from Werner, F. 18%. Beitrige zur Kentniss der Reptilien und Batnchier Guenero. Univ. Kansas Sci Bull. 54201-212. vonCenvllamerika undChile, sowieeinigerseltenerer Schlang- Hutweg, N. and J.A. Oliver. 1940. A contribution to the herpetology enarten Verh. 2.001. Bot. Ges. Win 46:344365. of the Isthmus of Tehuantepec. IV.A n annotatedchecklist of the Wilson, LD. and J.R Meyer. 1985. The snakes of Honduras. 2nd ed. amphibians and reptiles collected on the Pacific slope during Milwaukee Publ. Mus. thesummerof 1936. Misc.Pub1. Mus. k l .U niv. Michigan47:l- Woodbury, A.M. and D.M. Woodbury. 1944. Notes on Mexican 31. snakes from Ouaa.J . Washington Acad. Sci. 34:36&373. Lawson, R. and H.C.D essauer. 1981. Electrophoretic evaluation of Wright, A.H. and A.H. Wright. 1957. Handbook of snakes of the Unit- the colubrid genus Ehpbe Fitzinger. Isozyme Bull. 1483. ed States and Canada. Vol. 1. Constock Publ. Assoc., Ithaca, Lee, J.C. 1980. An ecogeognphic analysis of the herpetofauna of the New York. Yucatln Penihsula. Univ. Kansas Mus. Nat. Hist. Misc. Publ. (67): 1-75. -- Robert M. Price, Depvunent of Biology, New York University, New Martin, P.S. 1955. Herpadogical records from the GodzF dhs re- York, NY 10003. f7 gion of southwestern Tarmulipas, Mexico. Copeia 1955:173 -11 9850.8 .A biogeography ofreptiles and amphibiii n the Gomez Primary editor for this account, Larry David Wilson. Farias region, Tmaulipas, Mexico. Misc. PuM. Mus. Zool. Univ. Published 15 October 1991 and Copyright 0 1991 by the Society for Michigan 101:l-102. the Study of Amphibi and Reptiles. Mehrtens,J .M. 1987. Living snakes of theworld in color. Sterling Publ.

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