RE- SQUAMATk SERPENTES: SENTICOLS,S. TIUASPIS Catalogue of American Amphibians and .

Price, R.M. 1991. Senticolis, S. hiaspis. Senticolis Dowling and Fries SmticoIisDowlingandFries,1987:202.Type-, SenlicoIis~ Cope 1866, by monotypy.

Content. A single species, hiaspis, is recognjzed

Definition. Smticolisis a medium-sized, elongate, anteriorly and posteriorly tapered, long-tailed colubrine (smDowling 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 30-35sof 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+19to 33+39+23,with 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 fiiof the organ is smooth. At the distal point of the smooth region 256-282 in females, with females tending to have 15.4-20.5scales 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 onthe distal half of the mean dimorphism varying 13.2-22.2scutes 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 3-5stiff 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 . The cross- hatched area indicates a region of intergradation. Diagoosls. Senticolis mybe 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 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- Senficolishas 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 heidentification 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, bedon 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 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.pis:Dowling and Fries, 1987:202.

Content. Three subspecies are recognized: h'aqis, iw- medius, and mutabilk

Deflnttion and Magnosis. See the generic account.

Descriptiom. The original description was provided by Cope(1866). ~chmidtandhdrews (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 reclassificationof the nenus (Dowlinn and Fries. 1987). A recent description is induded in Wilson and ~Ger(1985):

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~fromsoutheasternArizona 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 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 completelyevertedand 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%), ~ilnther(la%), of F. Waite Gibson. Powell (1980), Ballinger and Lynch (1983), and ~tebbii(1985).

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 similaritiesin scutellation,the 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,ranging fromthephen Forest inGuatemala southwardto Costa Rica on the Caribbean side of Central America. The specimen Figure 3. Smlicdis biaspis inkmnedius from the Chhahua designatedastheholotypeof 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~~:Werner, 1896247. Type-locality, 'Honduras.' 11- Coluber tria.pis Cope, 1866128. See species account. lustrated on pl. 6, additional type data unknown. Nahix hiaspis: Cope, 1887:71. Ekapbe hiaspis mutubilis: hart, 1948:68. Scolophis mutabilis: Bocoun, 1888:680 (part). Ehpbe hiaspis in&media.. Mertens, 1952b:93. Ekaphe hiaspis hiaspix Stuart 1948:69. Senticolic hiaspis mutabilk. Dowling and Fries, 1987:202. 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 sd,rounded, pale spot at or near the with an elongate centnl opening. The band is usually broken at the middle of the interparietal suture. A blotched pattern is retained into frontoparietal suture. The adults are uniform pale brown dordy. adulthood. Remark. The hdotype was mistakenly given as USNM 6735 in the original desaiption.

Literature Cited Pilyopbis intennediusBoeuger, 1883:148. Type-locality, 'Mexico,' restriaedto Hacienda El Sabiio. Michoadn, Mexico by Dowlinn Alvvez del Toro, M. md H.M. Smith. 19%. Notulae herpetologicae (1960). Holotype, endt ten be& Museum (SMF) 34575, a juvg chiapuhe I. Herpetologka 123-17. nile rmle from the colledion of Dr. Pa~enSte~her..date 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 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 we, 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 cbbmmStone, 1911:231. Ehphemutabilis:Taylor, 1%@459.Fit useofthis 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 characterizedby 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& trfuspfsmutabUls (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.VertebratafromtheYuatan. Reptilia, bergixhen Naturf. Ges. 487:l-120. Amphibia, and Pisces. Bull. Mus. Comp. Zool. W146-159. -1952b. 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