576.1 REPTII& SQUAMATA SERPENTES: BOIDAE CORAILUS ENYDRIS
Catalogue of American Amphibians and Reptiles. hourglass shapes or round-cornered rhomboid-like shapes (Henderson, 1991: Pig. 1, paern 3). The venter is usually heavily Henderson, RW. 1993. Comllus enydh patterned with dark brown. (2) Grenada Bank: Maximum SVL is at least 1625 mm. On Grenada the dorsal ground color is extremely CoralZus enydris (Linnaeus) variable, including shades of yellow, orange, red, gray, taupe, brown (to nearly black), and olive. For snakes in which the dorsal ground Boa Envd* Linnaeus. 1758:215. Tvw-localitv. uAmerica." Hole color is vellowish. the dorsum mav be comoletelv devoid of att tern. ty&, ~aturhistorika~iksmusee( ~tockhoi&(~ndersson, 1899), lightly patterned aith flecks of brown that &cupy less than singld mssiblv uncatalonued, collector and date of collection un- scale. or have a Danern of dark flecks ~rovidin~the indication of an known[sex unknok, sVL 1020 mm (not examined by author). hourglass shape: Snakes that have a dorsal gro"und color other than Comllus en$&: Forcan, 1951:197. yellow exhibit tremendous variability in pattern shape (anteroposteriorlyelongated,dorsoventrally broadenedrhomb, hour- Conten+ Two subspecies are recognized, C. e. enydris and glass, spade; Henderson, 1991: Fig. 1, patterns 16). Dingy yellow is C. e. cooki. the predominant ventral ground color, but occasionally the venter is cream or white. The ventral pattern may be almost immaculate, Defhitlon MaximumSVLisat least 1870 mm(fromTrinidad). marked with flecks and spots, large blotches, or nearly covered with The general habitus is with a large chunky head, long anterior dark brown. Theventral panern usually becomesdenser posteriorly. maxillary teeth, slender neck, strongly laterally compressed body, On the islands of the Grenadines, yellow snakes have been collected and prehensile tail. CwaUus arydris exhibits virtually no sexual on Bequia and Union. Elsewhere dorsal ground color is gray, taupe, dimdrphism in pertinent scale chkcters. ~an~esin meritic features some shade of brown, or bright red-range (Mayreau only). The are as follows: dorsal scale rows at midbodv 37-63: ventrals 250-294: dorsal pattern is variable, but usually consists of some rhomboid subcaudals 94-133; ventrals + subcaudals 351-436;iupralabials (with derivative, spade, or hourglass shape (Henderson, 1991: Fig 1). 0) well-developed labial pits) 8-14; infralabials (with well-developed Costa Ria, Panad (and associated islands), and nonhern Colombia labial pits) 11-17; scales between supraorbitals 3-14; loreals 1-4; (north of the Cordillera Oriental): Maximum SVL is at least 1665 mm. subloreals 0-9; loreals + subloreals 1-11; circumorbital series 9-17; Dorsal ground color is yellowish, taupe, gray or some shade of brown nasals usually in contact with 4-14 scales bordering nasals. (browns predominate). The dorsal pattern is usually a series of lame Few snake species show greater color panern variation than C. rhombs benderson 1991: Fig. 1,- pattern 5). snakes from tks enydtis. Following are color pattern descriptions and notations of geographic area show more consistencythan elsewhere in the shape size from fwe geographic areas (some very broad) that describesome of the main dorsal element. The ventral ground color usually is dingy ofthe variability in this species- (1) St. Vincent: Maximum SVL is at yellow, and may be nearly immaculate to heavily patterned (if least 1374 mm. Dorsal ground color is gray, taupe, or brown; always patterned, more so on the posterior portion of the body). (4) with a conspicuous dorsal (- lateral because of strong lateral com- Trinidad, Tobago, lsla Margarita, and nonhern Venezuela: Snakes pression) pattern, the main elements of which are gray or brown from this area tend to be very large (especially fromTrinidad and the
Map. Distribution of Comllus enydris. The type-localities of both subspecies are too imprecise to plot. Solid circles mark known locality records. Subspecies of specimens from areas indicated by cross-hatching are undetermined. Orinoco Delta) and maximum SVL is at least 1870 mm. The dorsal principal nerves innervating the heat sensitive scales in Bullock and ground color is yellow, yellow-brown, khaki-brown, or copper Barren (1968). brown. Adistina dorsal panern may be absent, faint or conspicuous rhombs may be present, or the free edges of the dorsal scales may be Distribution. Corallus enydris has an extensive distribution dark brown. The venter may be whitish, cream, dingy yellow, or in the Neotropics, occurring from southwestern Costa Rica (south of bright yellow, and may be immaculate to heavily marked with dark 10°N) through Panad(including Isla del Rey, Isla Contadora, Isla de brown to black (becoming more predominant posteriorly). (5) Cebaco, Isla Suscantupu) into South America, where the species may Amazonia (including Atlantic coastal forest) and Guianan South be found in Colombia (east of the Andes, although Perez-Santos and America: Maximum SVL is at least 1640 mm. Dorsal ground color is Moreno [I9881 note specimens from west of the Andes). Venezuela yellow, some shade of brown or gray, but taupe is the predominant (including Isla ~arg&ta)andthrou~h~u~ana,Surinamejand~rench color. A dorsal pattern may be absent (if the dorsal ground color is Guiana into Amazonian Ecuador. Peru. Bolivia, and Brasil. In Brasil. yellow), but usually consists of a dorsovenmlly elongated rhomboi- C. enydris is encountered south of the-~ro~icbf~a~ricorn to 24043' dal shape (Henderson, 1991: Fig. 1, pattern 2), distinct or not, gray, in the state of SoPaulo, and also on Ilha Grande off the southeastern taupe, or brown in color. The venter may be white, cream, or coast. Insular distribution includes Trinidad and Tobago, and in the (predominantly) dingy yellow; immaculate, with a few flecks, scat- West Indies the species is known from St. Vincent, many of the tered spots and blotches, or nearly completely covered with dark Grenadines (Bequia, Ile Quane, Baliceaux, Mustique, Canouan, brown. The ventral pattern becomes heavier posteriorly. Mayreau, Union, Carriacou, Petit Maninique), and Grenada. Altitudi- The hemipenis in a C. e. enydris (from Venezuela south of the nal distribution is from sea level to about 1000 m, although Perez- KO Orinoco) is 19 mm long and undivided, the arms are reduced to Santos and Moreno (1988) note a specimen collected at 1900 m in terminal lateral bulges; the sulcus divides 10 mm from the base and Colombia. the forks run centrifugally to the tips of the lateral bulges; sulcal folds The distribution of C. e. enydris is largely coincident with the are large, slightly raised and unadorned; ornamentation is flounced; distributionof lowland tropical rainforest. These snakes are virtually the proximal third of the organ is nude, followed distally by three absent fromcaatinga in eastern Brasil, and, in general, fromareas that shallow, transverse flounces that fuse with the sulcal folds and receive habitats (from cactus and thorn scrub to is covered with scattered, shallow, rounded papillae (Branch, 1981). rainforest) and rainfall regimes, including areas that receive habitat (Sexton and Heatwole, 1965; enydris has 9-13subcaudals compared to 79-87 in C. annulatus. Roze, 1970; Cunha and Nascimento, 1978; Duellman, 1978, 1989, 65-74 in C. caninus, and 40in C. cropanii. Juvenile (yellow) C. 1m; Rivero-Blanco and Dixon, 1979; Henderson et al., 1979; caninus bear a superficial resemblance to yellow phase C. enydris, Hoogrnoed, 1979, Fugler, 1986, Henderson, 1988; Perez-Santos and but scale characters easily distinguish the two taxa. Moreno, 1988,1991;Henderson, 1990b, 1993a;Rodriguezand Cadle, 1990; Zimmerman and Rodrigues, 1990; Schwartz and Henderson, Descriptions. General descriptions are many and vary in 1991;Henderson and Winstel, 1992;Lillywhite and Henderson, 1993; usefulness; listed by country or area: Panad (Cope, 1876; Smith and Hendersonand Boos, in press), die1 activity(Cunha and Nascimento, Grant, 1958), Colombia (Dunn, 1944; Perez-Santos and Moreno, 1978; Henderson et al., 1979; Duellman, 1989,1990; Rodri~uezand 19881, Venezuela (Roze, 1964, 1966; Lancini V., 19791, Guyana Cadle, 1m;ZimmerrnanandRodrigues, 1930; ~endersonandwinstel, (Parker, 1935; Beebe, 1946), Suriname (Moonen et al., 1979), French 1992). defensive behavior (Battecki and Hevmarn. 1987: Greene. Guiana (Chippaux, 19861, Ecuador (Duellman, 1978; Perez-Santos lg88;'Kreutz, 1989), stereotyped behavior (Carpenter and Fergusonj andMoreno, 19911, Peru (CarrillodeEspinoza,1966,1970; Dionand 1977), diet (Urich, 1933; Wehekind, 1955; Smith and Grant, 1958; Soini, 1976,19861,Brasil Wagler, in Spu, 1824;Amaral,1976; Cunha SextonandHeatwole, 1965;Pendlebury, 1974;Cunhaand Naximento, and Nascimento, 19781,Trinidad and Tobago (Mole, 1924; Mole and 1978; Greene, 1983; Duellman, 1989,1990; O'Shea, 1989; Rodriguez Urich, 1894; Boos and Quesnel, 1969; Ernsley, 1977; Moonen, 1977; andcadle, 1990;Zimmerman and Rodrigues, 1990;Henderson, f91, Hardy, 1982; Hendersonand Boos, in press), West Indies (Henderson, 1993a. 1993b; Schwartz and Henderson, 1991; Lillywhite and 1988,1991). ~enderson,1993; Henderson and Boos, reiative abun- dance (Durn, 1949; Stemmler and Vesely, 1968; Roze, 1970; Fugler, Ilhrstrations. Many photographs of this geographically 1986), population density (Grenada) (Henderson and Winstel, 19921, widespread snake have been published. and it would serve no useful distribution of subspecies (Giinther, 1895; Parker, 1935; Lancini V., to attempt to list ali here. ~iackand white photographs 1979;Chippaux, 1986), phylogeny (Kluge, 19911, conservation strat- appear in Diunars (1942), Beebe (1946), Carrillo de Espinoza (1966), egy (Henderson, 1988), abuse by humans (Dunn, 13931, captive Boos and Quesnel(1%9), Foekema (19741, Cunha and Nascimento maintenance (Foekema, 1974; Stafford, 1986a, 1986b, 1986~;Abuys, (1978), Hardy (1982), Stafford (1986~1,Banecki and Heyrnarn (198i9, 1988; Winstel, 1989,1930,1992). Abuys (1988). Henderson (1988,1930a, 1990b), Winstel (1988,19901, Kreutz (1989), Oxtoby (1989), and Henderson and Winstel (1992). Etymology. The specific name enydris is from the Greek Color photographs are in Schmidt and Inger (1957), Roze (19701, meaning watersnake. The laterally compressed body may have Moonen et al. (1979), Lancini V. (1979), Manison (1986), Stafford suggested to Linnaeus that the species was adapted for an aquatic (1986), Mehnens(1987),Henderson(1988), Perez-Sgntos and Moreno existence. The name cooki is a patronym in honor of Edward Cooke, (l988,1991),Winstel (l988,1989,1992),Carnpbelland Lamar (1989), presumably the collector and/or donor of what became the type- and Ross and Marzec (1990). Black and white drawings are found in specimen. Wagler, in Spu (1824), Jan and Sordelli (18641, Roze (19661, Barren (1970), Emsley (1977), Stafford (1986b). Color drawings are in - Amaral(1976). Drawings of the hemipenis appear in Branch (1981). 1. CoraUus enydris enydrfs (Limaeus) Drawings of cranial elements occur in Underwood (1976) and Kluge (1931),drawingsofsoFttissue in Beddard(1908), anda diagramofthe Boa Enydris Linnaeus, 1758:215. See species synonymy. Figure 1. (top row lew. ComUus enydh from the vicinityof Layou, St. Patrick Parish, St. Vincent. Figure 2 (top row right). ComUus en## from he vicinity of Westerhall, St. David Parish, Grenada. Photognph by R.A. Winstel. Figure 3. (second row left). Comllus enydris from 1.0 mi NE Grand Etang, St. Andrew Parish, Grenada. Figure4. (second row right). ComUusenydrisfrom Hollis Resewoir, St. George Co., Trinidad. Photograph by Hans Eh. Boos. Figure 5. (third row left). Comllur enydris from 13 km S, 1 km E Puente Cuyuni, Bolivar, Venezuela. Photograph by W.E. Duellman. Figure 6. (thud row right). ComUusenydris from JurFi-Itatins, So Paulo, Bail. Photograph by I. Sazima. Figure 7. (fourth row left). Comlluseny.fris from C~ZcoAm;rZ~nica, Madre de Dios, Pem. Photograph by W.E. Duellman. BoaHortulana Linnaeus, 1758:215. Type-locality 'America." Holo- type, Naturhistoriska Riksmuseet, Stockholm (NRS.) Lin. 7, col- lector and date of collection unknown, sex and age class Abuys, A. 1988. De tuinboa (Comllusenydris) in literatuur, natuuren unknown (not examined by author). terrarium (1). Lacena 46:194-198. Vipem bitis Laurenti, 1768:102. Type-locality 'Brazil." Holotype, not Amaral, A. 1976. Serpentes do Brasil: iconografia colorada. Brazilian traced (Seba, 1735). snakes: a color iconography. Univ. SoPaulo, SoPaulo. Vipera madamis Laurenti, 1768:102. Type-locality, 'Isle Madere." Anderson, L.G. 1899.CatalogueofLinneantype-specimensofsnakes Holotype, not traced (Seba, 1734). in the Royal Museum in Stockholm. Handl. Svenska Vet. Akad. BoaMewemii Sentzen, 179653. Type-locality, unknown. Holotype, Bd. 24, Afd. 4(6):1-35. not traced. Barbour, T. 1914. Acontribution to the zoogeography ofthe West In- Boa Ambleocepbala Donndorff, 17983. Type-locality, not given. dies. with especial reference to amphibians and reptiles. Bull. Holotype, not traced. Mus. Comp. -%I. 44:209-359. Boaobtusiceps Bechstein, in Lace+&, 1802:46. Type-locality, none Barren, R 1970.The pit organsofsnakes, pp. 277-300. In C.Gansand given. Holotype, not traced. T.S. Parsons (eds.), Biology of the Reptilia. Vo1.2, Morphology Comllus obtusimrris Daudin, 1803:259. Substitute name for Boa B. Academic Press, New York. mmiiSentzen. Banecki, U. and E.W. Heyrnann. 1987. Piild observation of snake- Xipbo.com omaturn Wagler, in Spix, 1824:40. Type-locality 'in a- mobbing in a group of Saddle-back Tamarins, Saguinusfusci- quis flumini Solimoens confinibus." Holotype, Zoologische collis nigriJons. Folia Primatol. 48:199-202. Staatssammlung, Munich, 2694/0, collector and date of collec- Begak, W. 1965. Constituglo cromoss6mica e mecanismo de deter- tion unknown, sex unknown, subadult (not examined by au- minaglo do sexo em ofdios sul-Americanos. I. Aspectos cario- thor). tipicos. Mem. Inst. Butantan 3237-78. Xipko.com domale Wagler, in Spix 1824:43 (pl. XV). Substitute Beddard, F.E. 1908.Acomparison ofthe neotropical speciesof ComC name for Boa bortulana Ius, C. wokii, with C. madagascariensis:and on some points in Boa m&la Reuss, 1834:129. Type-locality, "Brasilien, Provinz II- the anatomvof Coralluscaninus. Proc. Zool. Soc. London 1908: heos." Holotype, Natur-Museums und Forschungs-lnstituts 135-158. Senckenberg, Frankfuna M. (SMF) 16810, collector, date of col- Beebe.. W. 1946.- Fieldnoteson the snakesof Kartabo. BritishGuiana. lection, sex, and agestatus unknown(notexamined by author). and Caripito, Venezuela. Zoologica 31:ll-51. Corallus maculatus Gray, 1842:42. Type-locality, 'Berbice." Holo- Boos, H.andV. Quesnel. 1969. ReptilesofTrinidad andTobago.Syn- type, British Museum (Natural History) (BMNH) 1946.1.10.43, color, Port-of-Spain. collector and date of collection unknown, female, 908 mm SVL Boulenger, GA. 1893. Catalogue of snakes in the British Museum (examined by author). (Natural History). Vol. I., containing the families Typhlopidae, Comllus bortulanus: Boulenger, 1893101. Glauconiidae, Boidae, Ilysiidae, Uropeltidae,Xenopeltidaeand Comllus enydris enydris: Forcart, 1951:197. First use of trinomial. Colubridaeaglyphae, part. British Museum (Nat. Hist.). London. Branch, W.R. 1981. Hemipenes of the Madagascan bogs Acranbpbis Diagnosis. Comllus e. enydris is best characterized by 50 or and Sanziniu, with a review of hemipeneal morphology in the more dorsal scale rows at midbody (rarely a Peruvian or Bolivian Boinae. J. Herpetol. 15:91-99. specimen will have 47 or 48); C. e. cooki invariably has Panama." Carrillo de Espinoza, N. 1966. Contribuci6n al conocimiento de 10s Holotype,Academy of Natural Sciences of Philadelphia (ANSP) boideos Peruanas (Boidae, Ophidia, Reptilia). Publ. Mus. Hist. 10325, collector, date of collection, sex, and age class unknown Nat. uJavier Prado", Ser. A., Zool. 21:86-136. (not examined by author). -.1970. Contribuci6n al conocirniento de 10s reptiles del Peru. Corallus cooki: Boulenger, 189399. Publ. Mus. Hist. Nat. "Javier Prado", Ser. A., %I. 221-64. Boagrenademu Barbour, 1914327. Type-locality, "St. George's, Chippaux,J.-P. 1986. Lesserpentsde la Guyane franpise.Coll. Faune Grenada." Holotype, MuseumofCornparativeZoology,Harvard Trop. 27:l-165. Univ. (MCZ) 7791, collected 20August 1910 byG.M. Allen, adult Cope, E.D. 1876. On the Batrachia and Reptilia of Costa Rica. J. Acad. male (examined by author). Nat. Sci. Philadelphia 893-154. Boa salmonidia Briceiio, 1934:1141. Type-locality, "Frontera con Cunha, 0.R da and F.P. doNascirnento. 1978. Ofidios da Amaz6nia. Colombia enel RiodeOro, y Distrito Col6ndel Edo Zulia." Hol- X As cobras da reg50 late doParii. Mus. Paraense Emilio Goel- otype, not traced. di, Publ. Avul. 31:l-196. Corallus enydris cookii: Forcan, 1951:197. First use of combination. Daudin, P.M. 1803. Histoirenaturelle,generaleet paniculieredesrep- tiles, Tome 5. Imprimerie F. Dufan, Paris. Diagnosis. Comllus e. cooki is best characterized by having Ditmars, RL. 1942. Snakes of the World. Macmillan Co., New York. <50 dorsal scale rows at rnidbody (as opposed to 50 or more in C.e. Dixon, J.R. and P. Soini. 1976. The reptiles of the upper Amazon Ba- enydris). The number of dorsal scale rows changes in a very short sin, Iquitos region, Peru. Contr. Biol. Geol. Milwaukee Pub. distance along the body of C. enydris. On the South American MUS.(12):l-91. mainland, the primary element in the dorsal pattern usually has -and -. 1986. The reptiles of the upper Amazon Basin, Iquitos rounded edges in C. e. a?ydris and a rhombic shape with sharper Region, Peru. Milwaukee Pub. Mus., Milwaukee, Wisconsin. edges in C. e. cook. Some specimens from the south bank of the Rio Donndorff, Jh. 1798. Zoologische Bepage zur XIII. Ausgabe des Orinoco in northeastern Venezuela exhibit a combination of < 50 LineischenNatursystems. Vol. 3. Wiedmamsche Buchandlung, midbodydorsal scale rowsand rounded pattern elementsat rnidbody. Leipzig. Snakes with <50 and >50 midbody dorsal scale rows occur in coastal Duellman, W.E. 1978. The biology of an equatorial herpetofauna in Guyana and Suriname,and possibly coastal French Guiana (Chippaux, Amazonian Ecuador. Univ. Kansas Mus. Nat. Hist. Misc. Publ. 1986). Although C. e. enydris exhibits mean differences ina number (65): 1-352. of characters (e.g., ventrals, number of scales between supraorbitals, -1989. Tropical herpetofauna communities: patterns ofcornmu- number of loreolabials), thesecharactersoverlap considerably, which nity structure in neotropical rainforests, pp. 61-88. In M.L. Har- makes them of limited diagnostic value. melin-Vivien and F. Bourliere (eds.), Vertebrates in complex tropical systems. Springer-Verlag, New York. Remark. The distribution of the subspecies of C. enydris in -. 1990. 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