Rlpituk SQUAMATA: SERPENTES: VIPERIDAE CROTALUS DURISSUS Catalogue of American Amphibians and Reptiles

Home , Caraiba, Crotalus durissus

RlPITUk SQUAMATA: SERPENTES: VIPERIDAE CROTALUS DURISSUS Catalogue of American Amphibians and Reptiles. body has 18-35 brownish or blackish diamond-shaped dorsal n blotches, usually with lighter centers, except in many individuals McCranie, J.R 1993. Cmtalus durissus. from southern South America in which the center is not noticeably lighter than the rest of the blotch. The blotches are usually outlined Crotalus durissus Linnaeus by one row of paler colored scales. Posteriorly, the blotches tend to Neotropical Rattlesnake become shorter, wider, paler in color, somewhat obscure, and merge with the primary lateral blotches. The primary lateral blotches lie Cmtalus Durissuc Linnaeus, 1758:214. Type-locality, -America," re- below the lateral points of the dorsal diamonds anteriorly. stricted tonJalapa,Veracruz, Mexico" by SmithandTaylor(1950: Paravertebral stripes that begin on the head posterior to and above 348), but not a good choice as the Veracruz population shows the eyes usually extend about 1-4 head lengths onto the neck. The some differences from those of the other populations of this paravertebral stripes can be poorly defined in some specimens from subspecies (see Klauber, 195260-61). Typespecimen original- northeastern Mexico or may extend less than one head lenmh onto ly in the Claudius Grill Suriname collection;later sent to the Zo- the neck in animals fromno&eastern and central Brasil. ~cu~ellation dlogiska Museet, Uppsala Universitet, Uppsala, Sweden, but is as follows: 25-33 scale rows at midbodv. all keeled exceot for the now lost (Klauber, 197234). See Remarks (1). lowest 1-3 rows on each side; 159-196 vktrals in males, 1&5-195in Cmtalopborur Durissus: Houttuyn, 1764313. females (C. uniwlor, if considered a subspecies of durissuc, would Umcmtalon Durirnrs: Fitzinger, 1843:29. lower the limits to 155 and 163, respectively); 22-34 subcaudals in Caudisona durissa: Cope, 1861:120.The first use ofthiscombination males, 18-31 in females; 11-18 supralabials; 12-20 infralabials. The that actually applied to this species. rostra1 usuallv is slinhtlv hinher than wide. The internasals and Caudbona durircuc: Sumichrast, 1881:270. prefrontals a& usuily 'paiGd and in contact medially, although Cmtalus akvissu: Richardson, 1972:lOl. Lapnu. snakes from the Pacific versant of Mexico northwest of the Isthmus of Cm&Jlusdu~:Jacobson, 1986:167. Lapnu,not seen,f&Smith Tehuantepec tend to have the prefrontals subdivided into smaller and Smith (1993:518). scales, as do occasional specimens from throughout the species range. Anteriorly 2-5 (usually 2 or 3) intersupraoculars are present. Content. Twelve subspecies are currently recognized: The parietal and occipital regions are covered by small keeled scales, durissus, ccatavella, collilineatus, culminatus, cumanensis, except in some specimens from northeastern Mkico in which larger, dtyina., mamfwnris, mnrimq tedJcus, tobnanrs, higonicus, irregular, vestigial parietals are present. The first supralabial usually and turbcan. See Remarks (2). is in broad contact with the prenasal, although a row of foveals may prevent this contact in specimens from southwestern Mexico. Each DeBnition and Magno8lr. Cmtalus durircuc is a large side of the head has 1-8 loreals (usually 2 or 3, except in snakes from species, with large adult males of some populations reaching about southwestern Mexico, which usually have 4 or 5). 1800 mm TL. However, several South American populations contain Cmtalus durissus is the only species of rattlesnake occurring members that are stunted, with many adults not exceeding about throughout much of its large geographical range. Only in Mkico 1000 mmTL. The species usually has a conspicuous vertebral ridge north of the Isthmus of Tehuantepec, in the savannas and alluvial that is best developed in large adults, especially on the anterior plains of the southwestern portions of the Venezuelan states of portion of the body. Individuals from northeastern Mkico tend to Anzoategui and Monagas, and on Aruba Island off the northern coast m have a ledweloped ridge. The dorsal ground color can be brown, of South America doesthe range of durissur overlap orapproachthat reddish brown, grayish brown, yellowish brown, yellowish gray, of other species of Cm&JIus (but see Remarks t21 concerning the yellowish olive, greenish gray, pale bluish gray, or nearly black. The South American 'species" C. uniwlor and C. qrandis). Cmtalus

Figwe. Cmtalus durirnrs totonacuc from pine-oak woodlands 5.6 km WSW El Lobo, Querhro, Mexico (photograph by L. Porras).

CL durissus approaches or overlaps the ranges of four large Mexican species (ahnr, basiliscur, momolaurrr, and scutulatus) north of the Isthmus of Tehuantepec. CmtaIus durissus can be distinguished Descriptions. An excellent morphological and color de- from each of these species by having conspicuous paravertebral scription of C. durissus is incampbell and Lamar (1989). Other good n stripes. Cmtalusdurisnrs can be further distinguished from C. abvx morphological and colordescriptionsare inCei(19861, Gloyd (1%0), and C. scutulatus by having a strongvertebral ridge (no strong ridge), Klauber (1952), Lancini and Kornacker (19891, Roze (19661, a nearly uniformly colored tail (strongly contrasting dark and pale Vanzolini et al. (1980), and Wilson and Meyer (1985). Osteological rings), and usually having 4-6, except 4-10 on the Pacific versant, features were described by Brattstrom (19641, and those of the scaie; in the inte&~-~kfro&l area (usually 8 or more). ~mtalk hemipenis by Klauber (1972) and Vellard (1946). durissus can be further distinrmished from C. basilism where their ranges overlap in southwestek Michoackn by having 27-33, usually .mwtrations. This is one of the most profusely illustrated 29, scale rowsat midbody (24-29, or usually 27), 170-182 and 173-188 species of all American amphibians and reptiles and the following ventrals in males and females, respectively (178201 and 184-206, citations are by no means exhaustive. Color illustrations of several respectively), and a nearly uniform tail color (gray with gray subspecies are in Campbell and Lamar (1989, Hoge (19661, and crossbands moderately evident). mica1 C. du- can be distin- Mehrtens (1987). Other good color illustrations are in Abuys (1987), guished from C. molossus by having a distinct vertebral ridge, in Alvarez del Tom (1983), Amaral (1977), Bolafios (19841, Campbell addition to having paravertebral stripes. However, individuals of and Brodie (1992), Carrizo (1988), Cei (1986), Chaves et al. (1990), durissus in northeastern Mexico, where the rangesofthe two species Coborn (19911, CNZ(1987), Duvernoy (1836-18491, Freiberg (19821, overlap, resemble C. rn0hw.s in tending to loose the distinct Freiberg and Walls (1984), Gans (19751, Hoge and Roman-Hoge vertebral ridge and in having poorlydefiied paravertebral stripes (198la), Jahn (19831, Klemmer (19631, Kundert (1974,19841, Lancini (Fig. 1). Campbell and Lamar (1989) have suggested that these two and Kornacker (1989), Levy (1983, as 'Fer-de-lance"), Mercado speciesmight hybridize innortheasternQuer&aro. ?he northeastern (1975), Moonen et al. (1979), Obst et al. (1988), Perez-Higareda and Mexican population of durisnrs can usually be distinguished from Smith (1991), Perez-Santos and Moreno (19881, Perkins (19741, Rob- mob in having paired parietal and occipital stripes (absent or erts (19841, RogC and Sauvanet (1987), Roze (19701, Schmidt (19901, obscure and ~oorlvdefied),a dark urefrontal crossbar (entire Schmidt and Inger (1957), Schnitzler (1987), and Villa et al. (1988). dorsum of headanterior to suboculars ukally dark), andthe primary Good black and white illustrations of several subspecies are in Glenn lateral blotches lvin~below the lateral wints of the dorsal diamonds and Straight (19821, Gloyd (1940), Harrisand Simmons (1972,197&), anteriorly (dokl bTotches more ofte; coalesced with the primary Hoge (1966), Hoge and Romano (19731, Hoge and Roman-Hoge lateral series). South American C.durhu canbe distinguished from (1981b), and Klauber (1956, 1972). Good black and whine illustra- C. vegmndis and C. unicolor by having well-defiied dorsal dii- tionsalsooccur inthe following papersthat arecitedelsewhere in this rnonds and paravertebral neck stripes (diamonds and stripes ob- account:Allenand Neil1 (1957),Amaral(1927b, 1977),~rrnstrongand scured by numerous scattered white-tipped scales in vegmndis and ~urphy(1979), Freiberg (19821, Gloyd and Kauffeld(19401, Harris diamonds and neck stripes faded and largely absent in adult and Simmons (1978b), Hoge and Romano-Hoge (1981a), Klauber Map 2. The SouthAmericandistribution of Cmtalurdurissuc. Open circlesmarktype-localities,solid circles otherrecords. The type-localities of the subspecies catcauella, collilinaahrs, and dryinas are too imprecise to be plotted (the type-locality restriction for catcauelka is invalid and that for dryinas is arbitrary). Some solid circles may represent several localities. ?he subspecific status of the many isolated Amazonian records is unknown. See McCranie (1984,1986) for the ranges of C. wgrandis and C. unicolor,both of which are considered by some authors to be subspecies of C. durissuc.

(1952), Lancini and Kornacker (19891, Roze (19661, Sage and em and central Michoadn, Mexico, to the western portion of the Capredoni (19711, Snchez-Herrera et al. (1981), Sandner Montilla Meseta Central in westcentral Costa Rica, with apparently continu- (1975, 19801, Stanek (19621, and Tinoco (1978). Other illustrations ous extensions into the Atlantic drainages of the Rio Grijalva depres- andtheirsubjectsare: drawings of headscalation (Abaloset al., 1964; sion in Chiapas, Mexico, and the middle and upper Rio Motagua Campbell and Lamar, 1989; Cei, 1986; Chippaux, 1987; Glenn and Valley in Guatemala. ?hi species also occurs in several isolated Straight, 19821, osteological features (Brattstrom, 1964; Carrizo, 1988; populations on the Atlantic versant. In Mtxico, populations occur in Duvernoy,1836-1849; Roze, 1970; Ruiz, 1952; Tinoco, 19781, west-central Nuevo Le6n, fromeast-central Tamaulipassouthwardto hemipenes (Prado, 1945; Roze, 1966; Vellard, 1946), karyotype extreme eastern Querttaro and northern Veracruz, in central (Becak, 1966; Gutierrez et al., 1979), microdermatoglyphic pattern Veracruz, and on the Peninsula de Yucat6n from its outer edge to its (Arroyo and Cerdas, 1986; Hoge and Romano-Hoge, 1983; Picado, base in northernGuatemala and Belize. In Guatemala, C. dunkws is 1931; Stille, 19871, aberrant coloration (Abalos and Nader, 1967; known from the upper Rio Negro Valley, and in Honduras from the Amaral, 1927a, 1927b, 1933a, 1934; Prado, 1945; Prado and Barros, Sula Plain and the Comayagua Valley. Although no records of this 1941; Renault and Schreiber, 19511, bicephalic specimen (Arnaral, species exist for Panama, Clark (1942, 1952) reported seeing rattles 1927~;Belluomini, Bisi et al., 1978), heterologous copulation from snakes from the dry tablelands around Tole, CaAazas, and La (Amaral, 1933b; Belluomini and Hoge, 1959). Mesa in the provinces of Chiriqui and Veraguas (also see Lyman, 1949). Summary information on distribution andlor signif~antlocal- Distribution In Middle America, Crotalur dutisms occurs ity records for the Middle American countries are Belize (Henderson more-or-less continuously along the Pacific versant from southwest- and Hoevers, 1975 [although these authors provide no documenta- tion fortheir overstated range of thisspecies in Belize, they docite the (1981), Wellborn et al. (1982); longevity in captivity, Snider and few previously published records for the country], 1977), Costa Rica Bowler (1992); feeding mechanism, Kardong et al. (1986); strike- (Savage and Villa, 1986; Taylor, 1951;Taylor et al., 19741, El Salvador induced chemosensory searching, Chiizar et al. (1978, 1982,19881, (FloresVilelaet al., 1991; Mertens, 19521, Guatemala (Campbell and Chiszar, Duvall et al. (1980), Duvall et al. (19801, Radcliffe et al. Vannini, 1989), Honduras (Cruz, 1987; Wilson and McCranie, 1991; (1980), Scudder et al. (1983); literature review on combat rituals, Wilson and Meyer, 1985), Mexico (Armstrong and Murphy, 1979; Carpenter (1986); reproduction, Araujo and Perazzolo (19741, Camarillo, 1983; Gloyd, 1940; Johnson, 1989; JuliP Zertuche, 1981; Dundee et al. (1986), Fitch (19851, Gongalves (19711, Langlada Klauber, 1952; Lee, 1980; Pbez-Higareda and Smith, 1991; Powell (1973), Langlada et al. (1974), Peterson (1983), Soldrzano and Cerdas and Parmerlee, 1980; Treviiio, 19801, and Nicaragua (Villa, 1984). (1988); birth defects, Langlada (1974); literature review on axial In South America, the species occurs in all mainland countries bifurcation, Smith and Perez-Higareda (1987); aberrant coloration, except Chile and Ecuador (but see below) and on the Venezuelan Hoge (1953), Villa (1984), also see Illustrations; parasites and/or islands of Margarita, Mom de la Iguana, and Tamarindo. Cmtalur diseases, Baker (1987) and Murphy and Armstrong (1978) reviewed unicolor,considered a subspecies of C. durissur by some herpetolo- much of the literature, papers not cited by them are Amyo et al. gists, occursonArubaIsland. InColombia, C. durissuc is known from (1980), Belluoimini, Saliba, and Abe (1978), Biasi et al. (1973,1976, the Rio Magdalena Valley, the llanos north of the Rio Guayabero/ 1981), Furlaneno et al. (19791, Johnson (1982), Machado (I%), Guaviare, and in the Caribbean lowlands from the Departamento de Moreno and Bolaiios (1977), Pesda and Biasi (19741, Pes& et al. Urdoba northeastward. In Venezuela, the species occurs through- (19741, Quinteroetal. (19901, Rego(1983a, 198313,1983~1,Santa Rosa out much of the northern and west-central portions, as well as in the et al. (1980); cytological and/or molecular studies, Becak (1966, Guiana highlands and a few isolated savannas in the southern part of 19681, Cadle (1992), Gutihrez et al. (1979), Miton (19921, Olmo the counuy. In the Guianas, a population occurs in the coastal (1984, l986), Prezoto et al. (lwl), Rodrifles and De Lucca (1973, lowlands from east-central Guyana to eastern French Guiana. Iso- 1974), Schreiber et al. (l%7), Sullivan (19741, Werrnan (1992); lated populations also occur in west-central and southwestern microdermatoglyphics, Arroyo and Cerdas (1986), Stille (1987); cra- Guyana. Northern Brazil has many isolated populations in AmapP, nial myology, Groombridge (1986), Haas (19731, Kardong et al. Pad, Roraima, Amazonas, and on Ilha de Maraj6. In eastern Brazil, (1986); vertebral meaurements, Christman (1975); sexual size differ- the species occurs more-or-less continuously from Cead and Rio ences, Fitch (1981); role in Mayan culture, Diaz-Bolio (1988); head Grande do None southward to the borders of Paraguay, Argentina, glands, Kochva (19781, Langlada et al. (19731, Lopes et al. (1972, and Uruguay, exclusive of the Atlantic coastal forests. In Bolivia, C. 1975); pit organ, Barren (1970); urinogenital system, Fox (1977); dut-issus occurs throughout most ofthe central portionofthe country lymphoid aggregation, Langlada et al. (1981); fine structure of liver east of the Andes with an extension into the Rio Madre de Dios cells, AlemPn (1972); coronary arteries, Erhart (1935); pulmonary drainage in southeastern Pen3. In Paraguay, the species occurs artery, Brongersma (1951); surgical techniques, Langlada and throughout most of the country except for the extreme northeastern Belluomini (1973a, 1973b), Langlada and Shinoiya (1973), Oliveira et portion. InArgentina,the speciesoccurseast ofthe Andessouthward al. (1988). A voluminous amount of literature is available on the to southern Mendoza andSanLuisprovinces. In Uruguay, C. durissur venom and snake-bite of this species. A good introduction to the occurs in the northern portion adjacent to Brazil and along the coast literature on the venom can be found in Ellion (19781, Mebs (19781, to the southeastern portion of the country in the Sierra de Mias near and the various chapters in Tu (1982) and that on snake-bite can be Solis de Mataojo. No actual records of C. durissur are known from found in Hardy (1989) and Watt (1989). Ecuador (Miyata, 1982; Peters, 1960), but the species may occur in southwestern Ecuador and adjacent Pen3 (Campbell and Lamar, Remarks. (1) Linaeus (1758) proposed the names bduc, 1989), as well as in the Departamento de HuPnuco, Pen3 (Schmidt and durissur, and dryinas for three species of rattlesnakes. The former m Walker, 1943). Summary information on distribution and/or signifi- two names were variously applied by early workers to the Neotropi- cant locality records for the South American countries are Argentina cal Rattlesnake and to the two North American species known today (Abalos et al., 1964; Cei, 1986; Freiberg, 1968; Sage and Capredoni, as C. borridur and C. adamanteus (see the synonymies for the 1971), Bolivia (Gloyd, 1940), Brazil (Cunha and Nascimento, 1980, respective species in Gloyd, 1940). Klauber (1941) provided a 1982; Gloyd, 1940; Hoge, 1966; Hoge and Romano-Hoge, 1981b; detailed analysis of the data and references provided by Linaeus Jorgeda Silva, 1993;MagalhBes, 1958; Miiller, 1968,1973; Nascimento (1758) and concluded that the name durisnrr was best applied to the et al., 1988; O'Shea, 19901, Colombia (Miiller, 1973; Perez-Santos and Central American populations of the Neotropical Rattlesnake, a Moreno, 1988; Renjifo, 1979), French Guiana (Chippaux, 1987; conclusion also reached by Gloyd (1940). Gloyd (1940) and Klauber Hoogmoed, 19831, Guyana (Allen and Neill, 1957; Gloyd, 1940; (1941) also pointed out that the South American populations that Harris and Simmons, 197813; Hoogmoed, 1983; Miiller, 1%8), Para- were being called Cmtalur tm+j?cus (Laurenti, 1768) were conspe- guay (Gallado, 1979; Gani, 1955; Gloyd, 1940; Scon and Loven, cific with the Central American populations. Unfortunately, some 19751, PerG (Carrillo de Espinoza, 1983; Meneses, 1974; Schmidt and South American workers have continued to use tanj?cus as the Walker, 19431, Suriname (Abuys, 1987; Hoge, 1964; Hoogmoed, specific name for this rattlesnake. Recently, Sandner Montilla (1975, 1983; Moonen et al., 19791, Uruguay (Achaval et al., 1978; Lema and 1983) even presented weak discussions vying to suppress the name FabiPn-Beurmann, 1977; Orejas-Mianda, 19691, and Venezuela durissuc in favor of tanj?cur (Hoge, 1966;Lanciniand Kornacker, 1989; Miiller, 1968,1973;Rivero (2) The South American subspecies of C. durissur are poorly Blanco and Dixon, 1979; Roze, 1966; Staton and Dion, 1977). defined and often difficult to distinguish. A thorough systematic Cmkaluc durisnrs is usually found in open, dry or semi-arid study of these populations, along with those occurring in Middle areas that have seasonal rainfall patterns. The species usually occurs America, is needed and was suggested by McCranie and Wilson below -700 m elevation, but occasional populations may occur as (1979) and Wison and Meyer (1985). The scalation data included in high as 2200 m in Middle America and 2500 m in South America. the subspecies definitions were taken from the literature. As a result, Campbell and Lamar (1989) have discussed the vegetation types and the ventral, subcaudal, and scale rows at midbody ranges given are altitudes where this species is found. considered deficient for many subspecies, especially those occurring in South America. Campbell and Lamar (1989) considered Cmkalur Fossil Record. Langebartel (1953) recorded "Cmtaluror vegtrandis Klauber and C. unicolw Van Lidth de Jeude to be Botbmpf vertebrae from early Holocene cave deposits in YucatPn, subspecies of C. dunbus. These were previously treated in the MQico. ~ataio~ueof American Amphibians and Reptiles as species separate from durissur byMcCranie(l984.1986). Subseauent~ublicationson Pertinent Literature. Klauber (1972) presented much these two taxa &e as follows: unicolor (color dhotos': Campbell and morph~lo~icaland biolo~icalinformation on this species. Campbell Lamar, 1989; Mehrtens, 1987; habitat photograph: Goode et al., 1990; andiamaAl989), ~mithand~mith(1976,1993), anhilla et al. (i988) fertility Loomis and Smith, 1987; microdermatoglyphics: Stille, 1987; listed much literature on this species, and Armstrong and Murphy species survival plan: Tryon, 1986; strike-induced chemosensory (1979) gave much informationon the natural history of the species in searching: Chiszar et al., 1992;Goode et al., 1990; studbook: Olney et MQico. Other works and their subjects are (with few exceptions, al., 1988; viral diseases: Smith et al., 1986), uegmndis (captive literature prior to 1970 is not cited here as it is usually cited in the reproduction: Muir, 1984; color photos: Campbell and Lamar, 1989; n papers mentioned): ecological notes within a snake community, Coborn, 1991; Kundert, 1984; Lancini and Kornacker, 1989; Henderson and Hoevers (1977), Via and Vangilder (1983); suggested Mehrtens, 1987; Tashjian, 1991; Trutnau, 1982; male courtship on a mimicry, Snchez-Herrera et al. (1981); behavioral notes in captivity, strike-induced chemosensory searching female: Chiszar et al., 1984; Chiszar, Wellborn et al. (1980), Cordeiro et al. (1981), Hoge et al. microdermatoglyphics: Stille, 1987; venom: Kaiser and Aird, 1987). Frost and Hillis (1990) suggested that Cmtalus durissus may be without providing any diagnostic characteristics. composed of several evolutionary species. Cmtalus durissus uucarella: Roberts, 1984:s. Lapsus. (3) Cmtaluspifanorum was described by Sandner Montilla (1980) on the basis of one, apparently adult, female specimen. The DeMtion. Paravertebral stripes on neck extend less than specimen was reported to be only652rnmTL. Otherthan itsapparent one head length, comprising 2-3 contiguous scale rows, without small size, the specimen falls within the range of variation reported distinctly lighter centers; short paraventral stripes are usually present for C. d. cumanensis and is tentatively included herein in the on neck; dorsal diamonds with distinctly lighter centers. Ventrals synonymy of that subspecies, as was suggested also by Campbell and number 167-175 in males, 172-180 in females; subcaudals 28-29 in Lamar (1989. males, 22-31 in females. Usually 4 scales are present in the internasal- (4) Cmtalus durisus neoleonensis Julia Zertuche, in Harris prefrontal area. Scale rows at midbody number 25-29 (usually 27 or and Simmons (1978a) isa nomen nudum, as no diagnostic character- 29). This subspecies is large, reaching at least 1600 mm TL. istics were stated nor was a holotype designated. Trevifio (1978) provided data from the three specimens upon which the name was 3. Crotalus durissus coUUineatus Amaral based (the number of ventrals given for one specimen is so low as to suggest a typographical error). From these data, this population Cmtalus tewi$cus var. collilineatus Amaral, 1927b:w (part). Type- appears very similar to C. d. totonacus. locality, 'zonas do Centro, S. E. e S.do Brasil e bem assirn da Ar- (5) Cmtalusdurissus trigonicus Harris and Simmons probably gentina, Paraguay e provavelmente Uruguay e Bolivii," restrict- is not distinct from C. d. ruruima, as variation within the few known ed to 'State Mato Grosso [Brasill" by lectotype selection (Hoge, specimens of hgonicus appears to be encompassed by that known 1966:139). Lectotype, Instituto Butantan Herpetologia 2180, for rumima. Abuys (1987), Cunha and Nascimento (19801, and sex, date, and collector not given (Hoge, 1%6:139) (not exam- O'Shea (1990) have also expressed some doubt as to the validity of ined by author). this taxon. In addition, problems associated with the holotype Cmtalus temicus cmtaminicus Goncalves, 1956:367. Type-locali- selection and the diagnosis provided by Harris and Simmons (1978b) ty, 'da regido de Morro Agudo, Franca e Ituverava [State of So are discussed by McCranie and Wilson (1979). Paulo, Brasill." No holotype designated. Nomen nudum, be- cause the description does not comply with the rules of the In- Etymology. The name durissw (L., durn - hard or tough; ternational Comrnision on Zoological Nomenclature. L., -issirnus - very) probably refers to the sturdy dorsal scales and Cmtalus durissus collilineatus: Hoge, 1%6:139. ridge in this species; uucavella is from the Portuguese word amtalusl. du?isu.d. collelineatur Langladaetal., 1974:253. Lapnrs. carcad, meaning rattle; wllilineatus (L., collum- neck; L., linea - Cmtalus durisnrs collineatus: Mercado, 197528. Lapsus. a line; L., -atus - provided with) refers to the paravertebral and paraventral stripes on the neck; culminatus (L., culmen- a ridge; L., DeMtion. Paravertebral stripes on neck extend more than -atus - provided with) refers to the strong vertebral ridge; dryinas is one head length, comprising 2-3 contiguous scale rows, usually probably derrived from the Greek word dryas - a wood or tree- without distinctly lighter centers; short paraventral stripes and acces- nymph, the meaning of which is unclear; marajoensis is named for sory blotches are usually present on neck; dorsal diamonds with Ilha de Marajo; ruruima is the local Indian name for Mount Roraima; distinctly lighter centers. Ventrals number 168-175 in males, 177-184 tempcus (L., temicus- dreadful or frightful) apparently refers to the in females; subcaudals 27-31 in males, 21-23 in females. Usually 4 fear invoked in humans by this snake; totonacus is named after the scales are present in the internasal-prefrontal area. Scale rows at Totonacs, an ancient Indian tribe of Veracmz, Mexico; trigonicus midbody number 25-31 (usually 27 or 29). This subspecies is large, (Gr., higonos - triangular; L., -icus - belonging to) apparently in reaching at least 1600 mm TL. reference to 'V shaped" markings on the supraoculars; tzabcan was said by the describer to be a Mayan word for rattlesnake. 4. Crotalus durissus culminatus Klauber

1. Crotalus durfsslcs durissus Llnnaeus Cmtalus durissur culminatus Klauber, 1952:65. Type-locality, 'El Sabino near Umapan, Michoadn, Mexico." Holotype, Field Cmtalus Durissw Linnaeus, 1758:214 (see species synonymy). Museum Natural History 126616 (formerly Edward H. Taylor Cmtalussimus Latreille, in Sonnini and Latreille, 1801:202. Type-lo- 52241, a juvenile female collected by E.H. Taylor and H.M. cality, 'Ceylan [in error]." No holotype designated, based on Smith in 1936 (not examined by author). Plate 45, fig. 4 in Seba (1735); also see Thireau (1991:4). Cmtalus tewiicus durkwr Amaral, 1929:5 (part). Defiaition Paravertebral stripes on neck extend more than Cmtalus durissur du-: Klauber, 1936:4 (part). one head length (1-3, usually about 2), comprising a single scale row Umtaluc]. tewiicus copmnus Amaral, 1937:162. vpe-locality, throughout more than half of their lengths and in 2 contiguousscale 'America Central." No holotype designated. Intendedasa sub- rows in remainder of length, without distinctlylighter centers; usually stitute name for Cmtalus durissw durissw. no paraventral stripeson neck; dorsal diamondswith distinctlylighter Cmtalusdutism durhs: Smith and Smith, 1993:518. Lapsus, origi- centers. Ventrals number 170-182 in males, 173-188 in females; nally used in Avila Soriano (1987:22), a dissertation. subcaudals 25-32 in males, 20-28 in females. Usually more than 4 scales are present in the internasal-prefrontal area. Scale rows at DeMtion. Paravertebral stripes on neck extend more than midbody number 27-33 (usually 29, less frequently 31). Thissubspe- one head length (1-4, usually 2-2 1/2), comprising 2-3 contiguous cies is large, reaching at least 1355 mm TL.. scale rows (paravertebral stripescan be lacking in some large adults), usually without conspicuously lighter centers; usually no paraventral 5. Crotalus durfssus cumanensis Humboldt stripes on neck; dorsal diamonds with somewhat lighter centers. Ventrals number 161-184 in males, 172-187 in females; subcaudals Cmtalus cumanensis Humboldt, 1813:6. Type-locality, 'Cumana 28-34 in males, 20-27 in females. Usually 4 scales are present in the Nenezuelal." No holotype designated. internasal-prefrontal area. Scale rows at midbody number 25-31 Cmtalus LoeJingii Humboldt, 18156. Type-locality, 'Cumana We- (usually 29, less frequently 27). This subspecies is large, reaching at nezuelal." No holotype designated. least 1800 rnm TL. See Remarks (2). Caudisona hjlingii: Cope, 1861:120. ~mtalusl.durissw cumanensis: Klauber, 1956:32 (as a possibly val- 2. Crotalus durfssus cascaveUa Wagler id form). Hoge (1966:142) formally recognized the subspecies and gave a definition of its purported range, but without pro- Cmtalus cascaoeNa Wagler, 1824:60. Type-locality, 'in campis providing any diagnostic characteristics. vinciae Bahiae." Hoge (1966:139) restricted the type locality to Cmtalus durisnrs eumanentis: Hoge,1967:223. Lapsus. "Mina Caraiba, Bahii [Brasill" by neotype selection (Instituto Cmtalus temifcus cumanensis: Sandner Montilla, 1975:51. n Butantan Herpetologia 23400), but Hoge's neotype selection is Cmtaluspifanorum Sandner Montilla, 1980:2. Type-locality, 'sesen- invalid (see Hoogmoed and Gmber, 1983). ta y ocho kil6metros al S. de la poblaci6n de Espino, en direc- Cmtalus tewi>cus var. wllirbombeatus Amaral, 1927b: 90. vpe-lo- ci6n de Puerto Parrnana, Parcelamiento de Agrotknicos 'Dr. cality, 'zona N. E. do Brasil." No holotype designated. Gonzalo Ledezma', entre los nacientes de 10s rios Otocuao 03- Umtal4. durissur carcatella: Klauber, 1956:32 (as a possibly valid te) y Carapa (Oeste), Distrito Infante, Estado GuPrico Nenezu- form). Hoge (1%6:139) formally recognized the subspecies elal." Holotype, Serpentario del Instituto Venezolano de Ofii- ologia CpNo. 1, anadult female, collected by HernPn Rauseo M, gradum elev. 45," restricted to uJuliode Castilho, MunicIpio Ta- in "a mediados" of 1980. See Remarks (3). quari, State Rio Grande do Sul, Brasil" by neotype selection Cmtalus durissus cuminamensis: Biicherl, 1980:122. Lapsus. (Hoge, 1%6:147). Neotype, Instituto Butantan Herpetologia Clmtalusl. durisnrsl.pifanotum: Pefaur, 192:15. 22997, an adult female, collector and date not given (Hoge, 1%6:147) (not examined by author). Deffnltloa Paravertebral stripes on neck extend more than Cmtalus scalbidus Boddaert, 1783:16. Type-locality not given. No one head length, comprising 2-3 contiguous scale rows, without holotype designated. distinctly lighter centers; paraventral stripes on neck long or consist- Cmtalus immaculatus Latreille, in Sonnini and Latreille, 1802:201. ing of many shorter broken stripes and usually more than one scale Type.-locality, "Indes orientales [in error]." No holotype desig- row wide; dorsal diamonds with distinctly lighter centers. Ventrals nated, based on Plate 95, fig. 3 and Plate %, fig. 1in Seba (17351, number 161-188 in males, 165-188 in females; subcaudals 22-34 in both plates reproduced by Aramata (1990:258 [the patternless males, 20-32 in females. Usually 4 scales are present in the internasal- snake] and 259, respectively); also see Thireau (191:2-3). prefrontal area. Scale rows at midbody number 25-31 (usually 27 or Cmtalus stmpitans Daudin, 1803:318. Type-locality, 'I'Amerique." 29). This subspecies is large, reaching at least 1750 mm TL. No holotype designated, based on Plate 95, fig. 3 and Plate 96, fig. 1in Seba (1735), both plates reproduced by Aramata (190: 6. Crotalus durfssus dtyfnas Linnaeus 258 [the patternless snake] and 259, respectively); also see Thireau (191:4). Cmtalus Dyinas Linnaeus, 1758:214. Type.-locality, "America," re- C[mtalus].temtcus: Cope, in Yarrow, 1875532 (part). stricted to "Paramaribo, Surinam" by Hoge (1966:143). Type- [Cmtalus&nijTcus var. &rn'j?cusl:Amaral, 1927b:90. By inference. specimen originally in the Museum Adolphi Friderici; later sent Cmtalus tenrims var. collilinaatus Amaral, 1927b:90 (part). See to the Zoologiska Museet, Uppsala Universitet, Uppsala, Swe- collilinmtus subspecies account. den, but now lost (Klauber, 1972:35). Paracmtalus temtcus: Reuss, 1930:88. Cmtalopbom Lhyinas: Houttuyn, 1764310. Cmtalus durissus femj?cus: Klauber, 1936:4 (part). Caudisona Drylnas: Laurenti, 176894. Cmtalus durisimus temtms: Stansk, 1962:63. Lapsus. Cmtalus stmpitans Var. dyinas: Daudin, 1803:320. Cmtalus durissus dyinus: Hoge, 1%6:142. Lapsus. Deffnltion. Paravertebral stripes on neck extend more than Cmtalus durissus dryinas: Klauber, 197235. one head length (usually at least three head lengths), usually comprising 2contiguous scale rows (paravertebral stripes can be indistinct in

Deffnltlon. Paravertebral stripes on neck extend more than some adults), without distinctly lighter centers;. long-* paraventral one head length, usually one scale row wide, without distinctly stripes on nedc (paraventral stribs can be indistinct in some adults); lighter centers; paraventral stripes on neck nearly continuous and dorsal diamonds without distinctlv lighter centers. Ventrals number one scale row wide; dorsal diamonds with distinctly lighter centers. 161-179 in males, 171-190 in femaiesrsubcaudals 27-32 in males, 19- Ventrals number 164-175 in males, 171-177 in females; subcaudals 25 in females. Usually 4scalesare present inthe internasal-prefrontal 24-32 in males, 23-24 in females. Usually 4 scales are present in the area. Scale rows at midbody number 25-33 (usually 27 or 29). This internasal-prefrontal area. Scale rows at midbody number 27-31. subspecies is large, reaching at least 1500 mm TL. This subspecies is large, reaching at least 1500 mm TL. 10. Crotalus durfssus totonacus 7. Crotalus durfssus marajoensfs Hoge Gloyd and Kauffeld

Cmtalusdurissur mamjoensis Hoge, 1966:143. Type-locality, "Tui- Cmtalustotonacus Gloyd and Kauffeld, 1940:12. Type-locality, 'Pa- uiu, llha de ~araj6,State par& Brasil." ~olotfpe,Instituto Bu- naco Island, about 75 miles south of Tampico, Veracruz, 12 tantan Herpetologia 17779, an adult female collected by A.R. miles inland fromCabo Rojo." Holotype, Chicago Academy Sci- Hoge, date-not g$en (not examined by author). ences 4469, a subadult female collected by Willis Woolems in May 1933 (not examined by author). DeMtioa Paravertebral stripes on neck extend more than Cmtaluc durissur totonacus: Smith and Taylor, 1945:190. one head length, comprising 2-3 contiguous scale rows, without Cmtalus basiliscus totonacus: Taylor, 1950:453. distinctly lighter centers; no paraventral stripes on neck (only a series Cmtalus durissus totonocus: Tu, 1971:91. Lapsus. of scattered black scales); dorsal diamonds with distinctly lighter Cmtalus durissus neoleonensis Juli5-Zertuche, in Harris and Sim- centers. Ventrals number 159-168 in males, 165-174 in females; mons, 1978a:lll. Nomen nu&. See Remarks (4). subcaudals 25-30 in males, 20-22 in females. Usually 4 scales are Clmtalusl. durisnrs totonacis: Gopalakrishnakone et al., 1979:57. present in the internasal-prefrontal area. Scale rows at midbody Lapsus. number 27-31 (usually 27 or 29). This subspecies is large, reachig at least 1500 mm TL. Deffnltion. Paravertebral stripes on neck usually poorly defined, with irregular borders; usually no paraventral stripes on 8. Crotalus durfssus rurufma Hoge neck; dorsal diamonds with somewhat lighter centers. Ventrals number 187-196 in males, 189-195 in females; subcaudals 24-29 in Cmtalus durissus mmima Hoge, 1%6:145. m-locality, 'Paulo males, 22-26 in females. Usually 4 scales are present in the internasal- Camp, 4000 ft., at Venezuelanversant of Mount Roraima, Vene- prefrontal area. Scale rows at midbody number 25-27. This subspe- zuela." Holotype, American Museum Natural History 36056, an cies is large, reachig at least 1800 mm TL. adult female collected by the Tate expedition in 1927 (not examined by author). 11. Crotalus durfssus trfgonfcus Cmtalus tenricus mmima: Sandner Montilla, 1980:7. Harris and Simmons Cmtalus durissus turima: Harding and Welch, 1980:66. Lapsus. Cmtalusduriwushigonicus Harrisand Simmons, 197&:112. Nomen Deffnltioa Paravertebral stripes on neck extend more than nudum. one head length, usually comprising 34contiguous scale rows, with Cmtalusdurissurtrigonicus Harris and Simmons, 1978b:305. Type- distinctly lighter centers; paraventral stripes or a series of dashes locality, "Rupununi Savanna of southwestern Guyana." Holo- usually present on neck; dorsal diamonds with distinctly lighter type, Natural History Society of Maryland RS 907 HSH/RSS, an centers. Ventrals number 167-172 in males, 174-177 in females; adult female, collector not given, collected in December 1970 subcaudals 25-32 in males, 21-23 in females. Usually 4 scales are (not examined by author). See Remarks (5). present in the internasal-prefrontal area. Scale rows at midbody number 27-29. Many specimens of this subspecies are stunted and Definition. Paravertebral stripes on neck extend more than ~1000mm TL, though some specimens do exceed 1500 mm TL. one head length, usually comprising 2 contiguous scale rows, without distinctly lighter centers; paraventral stripes or a series of dashes - 9. Crotalus durfssus t-icus (Laurenti) usually present on neck; dorsal diamonds with distinctly lighter centers. Ventrals number 170-172 in males, 177-178 in females; Caudisona &m>ca Laurenti, 1768:93. Type.-locality,'America infra subcaudals 29-31 in males, 21-23 in females. Usually 4 scales are present in the internasal-prefrontal area. Scale rows at midbody Iheringia, Zool. (45):5566. number 29-31. This subspecies is stunted, reaching about 1150 mm Armstrong, B.L. and J.B. Murphy. 1979. The natural history of Mexi- TL. can ranlesnakes. Univ. Kansas Mus. Nat. Hist. Spec. Publ. (5):vii + 88p. 12. CrotaZus durissus tzabcan Klauber Arroyo, O., R. Bolatios, and G. Munoz. 1980. The bacterial flora ofve- nomsand mouth cavities of Costa Rican snakes. Bull. Pan Arner. Cmtalus durissus tzabcan Klauber, 1952:71. Type-locality, "Kantu- Hlth. Org. 14:280-285. nil, Yucaan, Mexico." Holotype, Field Museum Natural History -and L. Cerdas. 1986. Microestructura de las escamasdorsalesde 36168, a young male collected by E. Wyllys Andrews on 28 Oc- nueve especies de serpientes costarricenses (Viperidae). Rev. tober 1939 (not examined by author). Biol. Trop. 34123-126. Cmtalus durisur tzabcan: Maruska, 1979:21. Lapsus,not seen, fde Avila Soriano, A. 1987. Algunos aspectos etnoherpetol6gicos de un Smith and Smith (1993:518). municipio totonaco de la Sierra Norte de Puebla: Tepango de Rodriguez. Los Reyes Iztacala, Tlalnepantla, Mkico. Tesis Bi- Demtion. Paravertebral stripes on neck extend from 1- ologo, Esc. Nac. Estud. Profesion. Iztacala, lalnepantla, Mexi- 2 1/2 (average about 11/21 head lengths, comprising 1-3 (2 predomi- co. nating, narrowing posteriorly) contiguous scale rows, without dii- Baker, M.R 1987. Synopsisof the Nernatoda parasitic in amphibians tinctly lighter centers; no paraventral stripes or spots on neck; dorsal and reptiles. Memorial Univ. Newfoundland Occ. Pap. Biol. diamonds with distinctly lighter centers. Ventrals number 177-188 in (1l):l-325. males, 182-191 in females; subcaudals 25-33 in males, 23-27 in Barren, R. 1970. The pit organs of snakes, p. 277-300. In C. Gans and females. Usually 4 scales are present in the internasal-prefrontalarea. T.S. Parsons (eds.), Biology of the Reptili. Vol. 2. Morphology Scale rows at midbody number 27-29 (more frequently 29). This B. Academic Press, London. subspecies is large, reaching at least 1750 mm TL. Becak, W. 1966 [19651. Constituiclo cromossBmica e mecanismo de determinaglo do sexo em ofidios Sul-Americanos. I. Aspect- Acknowledgments. I thank K. Adler. R.W. McDiarrnid. T.E. cariotipicos. Mem. Inst. Butantan 3237-78. Simmons, and H.M. Lithfor their help in obtaining copies of several -1968. Karyotypes, sex chromosomes, and chromosomal evolu- publications. Adler also provided information on Seba's rattlesnake tion in snakes, p. 5395. In W. Biicherl, E.E. Buckley, and V. plates. Deulofeu (eds.), Venomous animals and their venoms. Vol. I. Venomous vertebrates. Academic Press, New York. Literature Cited Belluomini, HA., P. de Biasi, G. Puorto, and V. Borelli. 1978 [1976/ 19773. Bicefalia em Cmtalusduriscus terriicus (Iaurenti). [Ser- Abalos, J.W., E.C. Baez, and R Nader. 1%4. Serpientes de Santiago pentes: Viperidae, Crotalinael. Mem. Inst. Butantan 40/41:117- del Estero (Republics Argentina). Acta Zool. Lilloana 20:211- 121. 283. -andA.R Hoge. 1959 [1957/19581. Um caso de c6pula heter6loga -and R Nader. 1967. Un caso de modificaci6n cromitica en Cm- entre Botbmps e Cmtalur (Serpentes). Mem. Inst. Butantan 28: talus durissus temicus. Rev. Fac. Cienc. Exac. Fisc. Nat., Univ. 91-94. de C6rdoba 28:l-12. -,A.M. Saliba,andA.S. Abe. 1978[1976/19773. Inqutritoanitomo- Abuys, A. 1987. The snakes of Surinam, Part XIX: family Viperidae, patoldgico em serpentes dos generos Cmtalus e Botbmpr [Ser- subfamily Crotalinae (Genus Cmtalus).Lineratura Serpentium, pentes, Viperidae, Crotalinae]. Mem. Inst. Butantan 40/41:123- English Ed. 7282-296. 128. Achaval, F., A.R. Melgarejo, and M. Meneghel. 1978. Ofidios del area Biasi, P. de, S.B. Pessaa, and H.E. Belluomini. 1973 [19721. Novasob- de influencia de Salto Grande (Aspectos biologicos y referen- sewagdes s6bre transmisslo congentia de hematozoirios de cias sobre ofidismo). V Reuni6n sobre Aspectos de Desarrollo serpentes peconhentas viviparas. Mern. Inst. Butantan 36:245- Ambiental, Cordoba. 249. Aledn,C. 1972. Notassobre la ultrastructura del higadoen tres espe- --, , G. Puorto, and W. Femandes. 1976 [19751. Nota sobre cies de serpientes. Mem. Soc. Cienc. Nat. La Salle 32:13&145. formas evolutivas de Trypanaroma de serpentes em meio de Allen, E.R and W.T. Neill. 1957. Some interesting rattlesnakes from cultura. Mem. Inst. Butantan 3985-101. southern British Guiana. Herpetologica 1367-74. -,E. de Riizo, P. Artigas, O.KA. Balderrama, and R.P. Cardoso,Jr. Alvarezdel Toro, M. 1983 [19821. Los reptiles de Chiapas. Tercera edi- 1981. 'In vitro" development of Trypanosoma butantanense ci6n, corregida y aumentada. Publ. Inst. Hist. Nat., Tuxtla Gu- Arantes and Fonseca, 1931. Rev. Inst. Med. Trop., SoPaulo 23: tirrez. 41-47. Amaral, A. do. 1927a (1926). Da occorrencia de albinism0 em casca- Boddaen, P. 1783. Specimen novae methodi distinguendi serpentia. vel, Cmtalus terrificus (Iaur.). Collectanea Ophiologica 6. Rev. Nova Acta. Phys. Med. Acad. Caes. Leop. Carol. 7:12-27. Mus. Paulista 15:55-57. Bolaiios, R 1984. Serpientes, venenos y ofidismo en Centroamtrica. -1927b (1926). VariagBes das marcas dorsaes de Cmtalus t&- Ed. Univ. Costa Rica, San Jose. ficus Laurenti, 1768. Collectanea Ophiologica 12. Rev. Mus. Brattstrom, B.H. 1964. Evolution of the pit vipers. Trans. San Diego Paulista 1589-91. Soc. Nat. Hist. 15185-268. -1927c (1926). Bicephalia em ophidios. Collectanea Ophiologi- Brongersma, L.D. 1951. Some remarks on the pulmonary artery in ca 13. Rev. Mus. Paulista 15:95-101. snakes with two lungs. 2001. Verh. (14):l-36. -1929. Key to the rattlesnakes of the genus Cmtalus Lint, 1758. Bucherl, W. 1980. Acdleos que matam. No mundo dos animais pe- Bull. Antivenin Inst. Amer. 54-5. conhentos. Livraria Kosmos Editoria, Rio de Janeiro, Brasil. -1933a [19321. Notassobre chromatismo de ophidiosI1. Casosde Cadle, J.E. 1992. Phylogenetic relationships among vipers: immuno- variaelo de colorido de cenas serpentes. Mem. Inst. Butantan logical evidence, p. 41-48. In JA. Campbell and E.D. Brodie, Jr. 7:81-87. (eds.), Biology of the pitvipers. Selva, Tyler, Texas. -19336 [19321. Contribui$%o&biologiadosophidiosdo Brasil IV. Camarillo, J.L. 1983. New herpetological records from the state of Sobre um caso de necrophilia heterologa na Jararaca (Botbmps Mkico. Bull. Maryland Herpetol. Soc. 1959-46. jararaca). Mem. Inst. ~utantan793-94. Campbell, J.A. and E.D. Brodie, Jr. (eds.). 1992. Biologyof the pitvi- -. 1934. Notas sobre chromatismo de ophidios 111. Um caso de pers. Selva, Tyler, Texas. xanthismo e umnovodealbinismo, observados no Brasil. Mern. -and W.W. Lamar. 1989. ThevenomousreptilesofLatinAmerica. Inst. Butantan 8:151-153. Comstock Publ. Assoc., Ithaca, New York. -1937 [1935-19361. Contribuiclo ao conhecimento dos ophidios -and J.P. Vannini. 1989. Distribution of amphibians and reptiles do Brasil VIII. Lita remissiva dos ophidios do Brasil. 2.a edicio. in Guatemala and Belize. Proc. West. Found. Verteb. Zool.4:l- Mern. Inst. Butantan 10:87-162, i-xix. 21. -. 1977. Serpentes do Brasil. Iconografia colorida. Brazilian Carpenter, C.C. 1986. An inventory of combat rituals in snakes. n snakes: a color iconography. Ed. Univ. de SoPaulo. Smithson. Herpetol. Info. Sew. (69):l-18. Aramata, H. 1990. Atlas anima. Volume 3. Amphibia and Reptilia (in Carrillo de Espinoza, N. 1983. Contribuci6n al conochiinto de las Japanese). Heibonsha Lmt., Publ., Tokyo. serpientes venenosas del Pen3 de las familias Viperidae, Elapi- Aradjo, M.L. de and M. Perazzolo. 1974. Nota prtvia sobre a biologia dae e Hydrophiidae (Ophidia: Reptilia). Publ. Mus. Hist. Nat. de tanatofidios em cativeiro. (Ophidia, Elapidae e Viperidae). 'Javier Prado", Ser. A, Zool. (30):l-55. Carrizo, G. 1988. La cascabel, 32 p. In Fauna Argentina. Anfibios y 163-436. In C. Gans and KA. Gans (eds.), Biology of the Rep reptiles, 2. Centro Ed. de America Latina, Buenos Aiies. tilia. Vol. 8. Physiology B. Academic Press, London. Cei, J.M. 1986. Reptiles del cenuo, cenuo-oeste y surde la Argentina. Erhart, M.B. 1935. The coronary cardiac arteries of snakes. Mem. Inst. Herpetofauna de las zonas Pridas y semlridas. Mus. Regionale Butantan 9:335-349. Sci. Nat., Torino. Monogr. (4):l-527. Pitch, H.S. 1981. Sexual size differences in reptiles. Univ. Kansas Mus. Chaves, F., J. Alvarado, R ~ynierich,and A. Solbrzano. 1990. Aspec- Nat. Hist. Misc. Publ. (70):l-72. - tos basicos sobre las serpientes de Costa Rica. Facul. Microbiol., -. 1985. Variation in clutch and litter size in New World reptiles. Inst. Clodomiro Picado, Univ. Costa Rica, San Jost. Univ. Kansas Mus. Nat. Hist. Misc. Publ. (76):l-76. Chippaux,J.-P. 1987 [1986l. Les serpentsde la Guyane frangaise. Coll. Fitzinger, L. 1843. Systema reptilium. Fasciculus primus. Amblyglos- Faune Trop. XXVII, &itions de I'ORSTOM, Paris. sae. Braumiiller et Seidel Bibliopolas, Vidobonae. Chiszar, D., J. Chiizar, T. Walsh, B. Demeter, and H.M. Smith. 1984. Flores Villela, OA., E. Hedndez Garcia, and A. Nieto Montes de Effect of male courtship on strike-induced chemosensory Oca. 1991. Catalogo de anfibios y reptiles del Museo de Zoolo- searching in a female ~rioanrattlesnake (Cmtalus uegmnd&) gIa, Facultad de Ciencias Universidad Nacional Aut6noma de at National Zoo. Bull. Marvland Hemetol. Soc. 20:145-151. Mhico. Ser. Cat. Mus. Zool. 'AlfonsoL.Herrera,"Cat. (3):l-222. -, D. Duvall, K. Scudder, and C.W. RaAcliffe. 1980. Simultaneous Fox, H. 1977.The urinogenital system of reptiles, p. 1-157. In C. Gans and successive discriminations between envenomated and andT.S. Parsons (eds.), Biology of the Reptilia. Vol. 6. Morphol- nonenvenomated mice by rattlesnakes (Cmtalus durissur and ogy E. Academic Press, London. C. tiridis). Behav. Neural Biol. 29:51&521. Freiberg, MA. 1968. Ofdios ponzoiiosos de la Argentina. Cienc. In- -,RK.K.Lee, C.W.Radcliffe, andH.M. Smith. 1992. Searching be- vest. 24:338353. haviors by rattlesnakes following predatory strikes, p. 369-382. -. 1982. Snakes of South America. T.F.H. Publ., Neptune, New In JA. Campbell and E.D. Brodie, Jr. (eds.), Biology of the pit- Jersey. vipers. Selva, Tyler, Texas. -and J.G. Walls. 1984. The world of venomous animals. T.F.H. -,P. Nelson, and H.M. Smith. 1988. Analysis of the behavioral se- Publ., Neptune, New Jersey. quence emitted by rattlesnakes during feeding episodes 111. Frost, D.R and D.M. Hillis. 1990. Species in concept and practice: Wie-induced chemosensory searching and location of rodent herpetological applications. Herpetologica 46:87-104. carcasses. Bull. Maryland Herpetol. Soc. 24:99-108. Furlanetto, R.S., H.E. Belluomini, H. Iizaka, and R. Rolim Rosa. 1979. -,C.W. Radcliffe, and H.M. Smith. 1978.Chemosensory searching Epizootia provocada por um bacilo difter6ide em serpentes forwounded prey by rattlesnakes is released by striking: a repli- mantidas em bioterio. Rev. Microbiol. 10:139-143. cation report. Herpetol. Rev. 954-56. Gallardo, J.M. 1979. Composici6n, distribuci6n y origen de la herpe- --, , and -. 1982. Predatory behavior of 'nervous" rattle- tofauna Chaqueiia, p. 299-307. In W.E. Duellman (ed.), The snakes, Cmtalusdurissurtemicus Herpetol. Rev. 13:90-91,93. South American herpetofauna: its origin, evolution, and disper- -, S. Wellborn, M.A. Wand, K.M. Scudder, and H.M. Smith. 1980. sal. Mus. Nat. Hist. Univ. Kansas Monogr. No. 7. Investigatory behavior in snakes 11: cage cleaning and the in- Gans, C. 1975. Reptiles of the world. Grosset & Dunlap Publ., New duction of defecation in snakes. An. Learn. Behav. 8505-510. York. Chriitman, S.P. 1975. The status of the extinct rattlesnake, Cmtalus Gatti, C. 1955. Las culebras venenosas del Paraguay. Rev. Med. Para- giganteus. Copeia 1975:43-47. guay 1:81-100. Clark, H.C. 1942.Venomoussnakes. Some Central American records. Glenn, J.L. and R.C. Straight. 1982. The rattlesnakes and their venom Incidenceofsnake-bite accidents. Amer. J.Trop. Med. 2237-49. yield and lethal toxicity, p. 3-119. In A.T. Tu (ed.). Rattlesnake -1952. Annual report of the Gorgas Memorial Laboratory, 1951. venoms. Their actions and treatment. Marcel Dekker, Inc., New q U.S. Gov. Print. Offfice, Washington, D.C. York. Coborn, J. 1991. The atlas of snakes of the world. T.F.H. Publ., Nep- Gloyd, H.K. 1940.The rattlesnakes, genera Sistmnrs and Cmtalus. A tune, New Jersey. study in zoogeography and evolution. Chicago Acad. Sci. Spec. Cope, E.D. 1861. An enumeration of the genera and species of rattle- Publ. (4):vii + 266 p. snakes, withsynonymy andreferences, p. 119-126. In S.W. Mit- -and C.F. Kauffeld. 1940. A new rattlesnake from Mexico. Bull. chell. Researches upon the venom of the rattlesnake: with an Chicago Acad. Sci. 6:ll-14. investigationof the anatomy and physiology of the organs con- Gongalves, J.M. 1956. Estudos &bre venenos de serpentes Brasilei- cerned. Smithson. Contr. Knowl. (12):viii + 145 p. ras. I1 - Cmtalus temtcus cmtaminicus, subesp6cie biolbgica. Cordeiro, C.L.S., A.R Hoge, and P. Sawaya. 1981. CriagBo de serpen- An. Acad. Brasileira Cien. 28:365-367. tes em cativeiro. Rev. BiotCrios 1:25.30. Gongalves, M.F. 1971. DeterminaGo da Cpoca de fecundidade em CNZ, GA. 1987. Serpientes venenosas de Honduras. Univ. Nac. Au- fernas do genero Cmtalur Arq. Mus. Nac., Rii de Janeiro 54: ton. Honduras, Tegucigalpa. 281-282. Cunha, 0.R da and F.P. do Nascimento. 1980. Ofidios da Amadnia Goode, M., C.W. Radcliffe, K. Estep, A. Odum, and D. Chiszar. 1990. XI - Ofidios de Roraima e notas sobre Erytbmlampnrs bauper- Fieldobservations on feediig behavior inanAruba Island rattle- tbuuii Dudril, Bibron & Dumeril, 1854, sin6nimo de Erytbm- snake (Cmtalus durisnrs unicolm): strike-induced chemosen- lampiusaescu@iiaesculapii (Linnaeus, 1758). Bol. Mus. Para- sory searching and trail following. Bull. Psychon. Soc. 28:312- ense Emilio Goeldi, Nova Serie, Zool. (102):l-21. 314. -and -. 1982. Ofidios da hazania XIV -As esp6cies de Mi- Gopalakrishnakone, P., B.J. Hawgood, R.D.G. Theakston, andA.H. nunrs, Botbmpq Lacbesue Cmtalus do sul do Par5 e oeste do Reid. 1979. An investigation of the specificity of antibodies MaranMo, incluindo Preas de cerrado deste estado. (Ophidia: raised to the crotoxin complex (from the venom of Cmtalus du- Elapidae e Viperidae). Bol. Mus. Paraense Emilio Goeldi, Nova rirruc tem~cus)using the Lethods of double-immunodiiusion Serie, 2.001. (112):l-58. and the enzvrne-linked immunosorbent assav (ELISA). Toxicon Daudin, F.M. 1803. Histoire naturelle gCnerale et particuli6re des reptiles. Vol. 5. F. Dufart, Paris. Groombridge, B. 1986. Comments on the M.p&tygoideurglandulae Diaz-Bolio, J. 1988. Why the rattlesnake in Mayan civilization. Pri- of crotaline snakes (Reptilia: Viperidae). Herpetologica 42:449- vately printed, MCrida, Yucatin, MCxico. 457. Dundee, HA., DA. White, and V. Riio-Gray. 1986. Observations on Gutieiiz,J.M., R.T. Taylor, and R Bolaiios. 1979. Cariotipos de diez the distribution and biology of some Yucatan Peninsula am- especies de serpientes costarricenses de la familia Viperidae. phibians and reptiles. Bull. Maryland Herpetol. Soc. 2237-50. Rev. Biol. Trop. 27309-319. Duvall, D., K. Scudder, and D. Chiszar. 1980. Rattlesnake predatory Haas, G. 1973. Muscles of the jaws and associated structures in the behaviour: mediation of prey discrimination and release of Rhynchocephalia and Squamata, p. 285-490. In C. GansandT.S. swallowing by cues arising from envenomated mice. An. Beh. Parsons (eds.), Biology of the Reptilia. Vol. 4. Morphology D. 28:674-683. Academic Press, London. Duvernoy, G.L. 1836-1849. Le Sgne animal distribue d'aprss son or- Harding, KA. and K.R.G. Welch. 1980. Venomous snakes of the ganisation, pour servir de base a I'histoire naturelle des ani- world. A checklist. Pergamon Press, Oxford, England. n maux, et d'introduaion a I'anatomie compa&e, par Georges Hardy, D.L. 1989.Annotated bibliography ofsnakebite inLatinher- Cuvier. Troisieme classe. Des animaux vertCbds. Les reptiles. ica, p. 14-17. In JA. Campbell and W.W. Lamar. Thevenomous Victor Masson, Paris. reptiles of Latin America. Comstock Publ. Assoc., Ithaca, New Elliott, W.B. 1978.Chemistryand immunologyofreptilianvenoms,p. York. Harris, H.S., Jr. and R.S. Simmons. 1972. Keys to the neotropical spe- -1952. Taxonomic studies of the rattlesnakes of mainland Mexi- cies and subspecies of the Cmtalus durissur group. Bull. Mary- co. Bull. Zool. Soc. San Diego (26):l-143. land Herpetol. Soc. 83-40. -19%. Rattlesnakes. Their habits, life histories, and influence on -and -. 1978a. A preliminary account of the rattlesnakes with mankind. Univ. California Press, Berkeley. Vol. 1. the descriptions (sic) of four new subspecies. Bull. Maryland -1972. Rattlesnakes. Their habits, life histories, and influence on Herpetol. Soc. 14:105-211. mankind. 2nd ed. 2 Vol. Univ. California Press, Berkeley. -and -. 1978b [1976/19773. A new subspecies of Cmtalus du- Klemmer, K. 1963. Lite der rezenten Giftschlangen. Elapidae, Hydro- rissuc (Serpentes: Crotalidae) from the Rupununisavanna (sic) pheidae, Viperidae und Crotalidae, p. 255-464. In Die Gift- of southwestern Guyana. Mem. Inst. Butantan 40/41:305-311. schlangen der Erde. Wukungen und Antigenitat der Gifte The- Henderson, R.W. and L.G. Hoevers. 1975. A checklist and key to the rapie vonGiftschlangenbiisen. N.G. E1wertUniv.-Verl.-Buchh., amphibians and reptiles of Belize, Central America. Milwaukee Marburghhn. Pub. Mus. Contr. Biol. Geol. (5):1-63. Kochva, E. 1978. Oralglands oftheReptilia, p. 43-161. In C.Gansand -and -. 1977. The seasonal incidence of snakes at a locality in KA. Cans (eds.), Biology of the Reptilii. Vol. 8. Physiology B. northern Belize. Copeia 1977349-355. Academic Press, London. Hoge, A.R 1953 [19521. Herpetologische notizen. Mem. Inst. Butan- Kundert, F. 1974. Fascination. Schlangen und Echsen. Serpents et It- tan 24269-270. zards. Snakes and lizards. F. Kundert, Spreitenbach. -1964 [1960/19621. Serpentes da fundag8o"Surinaam Museum." -. 1984. Das neue Schlangenbuch in Farbe. Albert Miiller Verl., Mem. Inst. Butantan 3051-64. Ziirich. -1966 [1%51. Preliminaryaccount on neotropical Crotalinae [Ser- Lancini, A.R and P.M. Kornacker. 1989. Die Schlangen von Venezu- pentes Viperidae]. Mem. Inst. Butantan 32:109-184. ela. Verl. Armitano Editores CA., Caracas. -1967. Serpentes do Territdrio Federal do ha$. Atas do Sim- Langebartel, DA. 1953. The reptiles and amphibians, p. 91-108. In p6sio s6bre a Biota AmazBnica 5 (Zool.):217-223. RT. Hatt, H.I. Fisher, Dh. Langebartel, and G.W. Brainerd. -,A. Rand, and PA. Federsoni, Jr. 1981. Manuten60 e criag8o de Faunal and archeological researches in Yucatan caves. Cran- serpentes em cativeiro. Rev. Biottrios 1:63-73. brook Inst. Sci., Bull. (33):l-119. -and SA. Romano. 1973[19721. Sinopsedas serpentespegonhen- Langlada, F.G. de. 1973 [19721. Ciclo sexual bienal de serpentes Cm- tasdo Brasil. Serpentes, ElapidaeeViperidae. Mem.Inst. Butan- talus do Brasil - Comprovafio. Mem. Inst. Butantan 3667-72. tan %:log-208. -1974 [19731. Anomalias congenitas em uma ninhada de 'casca- -and S.A.R.W.L. Romano-Hoge. 1981a [1978/19791. Poisonous veis." Mem. Inst. Butantan 37:239-251. snakesof the world. Part I. Checklist ofthe pitvipersViperoidea, -andH.E. Belluomini. 1973a [19721. Contribuigiioatecnica opera- Viperidae, Crotalinae. Mem.Inst. Butantan 42/43:179-310. t6ria de serpentes I. Hemipenicectomia bilateral em serpentes. -and -. 1981b [1978/19791. Sinopse das serpentes pegonhen- Mem. Inst. Butantan 36:73-78. tas do Brasil (2.a 4.). Mem. Inst. Butantan 42/43:373-496. -and-. 1973b [19721. Contribuifio atknicaoperat6riadeser- -and -. 1983. Notes on micro and ultrastructure of 'oberhaut- pentes 111. Ablagiio de gliindulas de veneno em serpentes do schen" in Viperoidea. Mem. Inst. Butantan 44/45:81-118. gCnero Cmtalus. Mem. Inst. Butantan 36:89-100. Hmgmoed, M.S. 1983 [19821. Snakes of the Guianan region. Mem. --, ,and J.C. Machado. 1973r19721. ConseqiiCnciasda ablagiio Inst. Butantan 46:219-254. cidrgica da gl5ndula principal de veneno em Cmtalus: Com- -and U. Gruber. 1983.Spix and Wagler type specimensof reptiles portamento do animal e estudo histopatol6gico da gliindula and amphibians in the Natural History Musea in Munich (Ger- acess6ria. Mem. Inst. Butantan %:101-108. many) and Leiden (The Netherlands). Spixiana, Suppl. @):319- -,M.F. Gongalves, and E.T. Rodrigues. 1974 [19731. Determinagso 415. da epoca de fecundidade em fCmeas do genera Cmtalus. Mem. Hounuyn, M. 1764.Natuurlyke historie of uitvoerige Beschryving der Inst. Butantan 37:253-260. Dieren, Planten en Mineraalen, Volgens het Samenstelvan den -,RL. Moraes, Jr., L. Denaro, and J.C. Machado. 1981[1978/1979]. Heer Limaeus. Vol. 6. Part 1. F. Hounuyn, Amsterdam. Sobre a possibilidade de um acirmulo linf6ide, encontrado em Humboldt, A. de. 1813. Sur deux nouvelles esp&cesde Crotales, p. 1- antro cloaca1 de serpentes, corresponder a bursa de fabricius 8. In A. Humboldt and A. Bonpland. Recueil d'observations de das aves. Mem. Inst. Butantan 42/43:367-371. zoologie et d'anatomie compafie faitesdans I'Octan Atlan- -and N. Shinoiya. 1973 [19721. Contribuigiio a acnica operat6ria tique, dans I'interieur du Nouveau Continent et dans la Mer du de serpentes 11. Derivafio intestinal, colostomia e cloacorrafii Sud pendant les amees 1799,1800,1801,1802et 1803. Vol. 2. (para obtengiio de urina sem contamina@o fecal em cloaca de F. Schoell and G. Dufour, Paris. serpentes). Mem. Inst. Butantan 3679-88. Jacobson, E.R 1986. Parasitic diseasesof reptiles, p. 162-181. In M.E. Laurenti,J.N. 1768. Specimenmedicum, exhibens synopsin reptilium Fowler (ed.), Zoo and wild animal medicine. Saunders, Phila- emendatam cum experimentis circa venena et antidota reptili- delphia. um austriacorum. Joan. Thomae Trattnern, Vinnae. Jahn, T. 1983. Brehms neue Tierenzyklopadie. Bd. 9. Reptilien, Am- Lee, J.C. 1980. An ecogeographic analysis of the herpetofauna of the phibien. Prisma Verlag, Giitersloh. Yucatan Peninsula. Univ. Kansas Mus. Nat. Hist. Misc. Publ. Johnson, J.D. 1989. A biogeographic analysis of the herpetofauna of northwestern Nuclear Central America. Milwaukee Pub. Mus. Contr. Biol. Geol. (76):1-66. nar dos fipteis da regiiio da fronteii Brasil-Uruguai. l&ringia, Johnson, J.W. 1982. A viral epizmtic in viperid snakes at Louisiana Zool.50:61-92. Purchase Gardens & Zoo, p. 106-107. In 4th annual reptile sym- Levy, C.K. 1983.A field guide to dangerous animals of North America posium on captive propagation & husbandry. 2001. Consor- including Central America. The Stephen Greene Press, Braale- tium, Inc., Thurmont, Maryland. boro, Vermont. Jorge da Siva, N., Jr. 1993. The snakes from Samuel Hydroelectric Limaeus, C. 1758. Systema naturae per regna tria naturae, secundum Power Plant and vicinity, Rondbnia, Brazil. Herpetol. Nat. Hist. classes, ordines, genera, species, cum characteribus, differem 137-86. tiis, synonymis, locis. Tomus I. Editio Decima, Reforrnata. Lau- Julil Zertuche, J. 1981. Reptiles mexicanos de importancia para la sa- rentii Salvii, Holmiae. lud pirblica y su distribuci6n geoefica. Salud Pirblica Mexico Loomis, M. and R. Smith. 1987. Fertility following caesarean section 23:329-343. inan ArubaIsland rattlesnake Ctutalusunicolor. Int. Zoo Yearb. Kaiser, 1.1. and S.D. Aird. 1987. Acrotoxin homolog from the venom 26:187-188. of the Uracoan rattlesnake (Cmtalus uegrandis). Toxicon 25: Lopes, RA., C. de Oliveira, G.M. Camps, and J.M. de Barns. 1975 1113-1120. [19741. Estudo morfol6gico e histoquimico da glPndula de Har- Kardong, K.V., P. Dullemeijer, and JAM. Fransen. 1986. Feeding der de alguns repteis Brasileiros I. Ophidea. Mem. Inst. Butan- mechanism in the rattlesnake Cmtalusdurissus Amph.-Rept. 7: tan 38:41-50. 271-302. -,V. Valeri, and C. de Oliveira. 1972. Histochemical study of mu- Klauber, L.M. 1936. Astatistical study ofthe rattlesnakes. I. -1ntroduc- copolysaccharides in the lacrymal gland of Cmtalusdutissus tion. Occ. Pap. San Diego Soc. Nat. Hist. (1):2-7. terrj?cus (Ophidia, Viperidae). AM. Histochem. 17:30>310. -1941. Four papers on the applications of statistical methods to Lyman, VA. 1949.Incidence of rattlesnakes in Panama. Herpetologi- herpetological problems. N. The rattlesnakes listed by Linnae- ca 544. us in 1758. Bull. Zool. Soc. San Diego (17):81-95. Machado,J.C. 1966 [19651. Formagbes cartilaginwas em midrdiode Cmtalus d tanfma Mem. Inst. Butantan 32:27-32. anPlisis taxon6mico y mogeogdfico. Ophidiofauna of Vera- MagalhSes, 0. de. 1958. Campanha antiofidica em Minas Gerais. cruz; taxonomical and zoogeographical analysis. Univ. Nac. Mem. Inst. Oswaldo Cruz 56:291-372. Aut6n. Mcbico, Pub. Esp. (7):l-122. Maruska, E.J. 1979. Animal census. Your Cincinnati Zoo News, 1979 Perez-Santos, C. and A.G. Moreno. 1988. Ofidios de Colombia. Mus. (summer):14-23. Reg. Sci. Nat., Torino, Monogr. (6):l-517. McCranie, J.R 1984. Cmtalus uegmndis. Cat. Amer. Amph. Rept.: Perkins, L. 1974. All color book of reptiles. Octopus Books, London. 350.1-350.2. Pesda, S.B. and P. de Biii. 1974 [19731. ConsideragBestaxon8micas -. 1986. Cmtalus unicolor. Cat. Amer. Amph. Rept.:389.1-389.2. sobre cistos esquizog8nicos e sobre gamet6citos de Hepatozo- -and L.D. Wilson. 1979. Commentary on taxonomic practice in on (Sporozoa, Haemogregarinidae)parasitas de serpentes Bra- regional herpetological publications: a review of Apreliminaty sileiras. Mem. Inst. Butantan 37:291-298. accountof tbe mttlesnaheswitbtbehcription offournewsub- --, ,andG. Puorto. 1974. Transferenciasdo Hepatom tupi- species by Herbert S. Hams, Jr. and Robert S. Simmons (1978. nambis, parasita de largerta Tupinambis teguiw'n, para a ser- ~ull.Mar$and Herpetol. s&,, 14(3):105-211), with comments pente cascavel (CmtalusdurirFuctem$cus), por intermMio do onother Hams and Simmons rattlesnake .uaDers. . Hemtol. Rev. mosquito Cukzfatigans. Mem. Inst. Oswaldo Cruz 72:295-299. 10:18-21. Peters, Jh. 1960. The snakes of Ecuador. A check list and key. Bull. Mebs, D. 1978. Pharamacology of reptilian venoms, p. 437-560. In C. Mus. Comp. Zool. 122:491-541. GansandKA. Gans (eds.), Biologyof the Reptilia. Vol. 8. Physi- Peterson, K.H. 1983. Reproduction of captive Cmtalus mitcbelli ology B. Academic Press, London. mitcbelli and Cmtalusdurisnrs at the Houston Zoological Gar- Mehrtens, J.M. 1987. Living snakesofthe wodd incolor. Sterling Publ. dens, p. 323-327. In D.L. Marcellini (ed.), Proceedings 6th rep Co., New York. tile symposium on captive propagation and husbandry. Zool. Meneses, 0.1974. Ofidios y ofdismo en el Perb. 11. 'Aspectos ecol6- Consortium, Inc., Thurmont, Maryland. gicos de la fauna ofidica ponzofiosa." Rev. Inst. Zoonosis E In- Picado, C. 1931. Epidermal microornaments of the crotalinae. Bull. vest. Pecuar. 2:79-84. Antivenin Inst. Amer. 4:104-105. Mercado, N.K. 1975. Ofidiosde Bolivia. Acad. Nac. Cienc., Bolivia. La Powell, R and J. Parmerlee. 1980. Geographical distribution: Cmtal- Paz. us durissur. Herpetol. Rev. 11:116. Mertens, R. 1952. Die Amphibien und Reptilien von El Salvador, auf Prado, A. 1945. Serpentes do Brasil. Ed. 'Sitios e Fazendas", SoPau- Grund der Reinvon R. Mertens und A. Zilch. Abh. Senck. Na- lo. turf. Ges. (487):l-83. -and F.P. de Barns. 1941 [19401. Notas ofiologicas 9. Duas cas- Minton, SA. 1992. Serologicrelationships among pitvipers: evidence caveis albinas do Brasil. Mem. Inst. Butantan 14:31-33. fromplasmaalbuminsand immunodiffusion, p. 155-161. In JA. Prezoto, B.C., E. Hiraichi, F.M.F. Abdalla, and Z.P. Picarelli. 1991.A~- Campbell and E.D. Brodie, Jr. (eds.), Biology of the pitvipers. tivation of the kallikrein-kinin system in plasma of some Brazil- Selva, Tyler, Texas. ian snakes. Comp. Biochem. Physiol. C, Comp. Pharmacol. Mita,K. 1982.Achecklistofthe amphibiansand reptilesof Ecuador Toxicol. 99:135-139. with a bibliographyof Ecuadorian herpetology. Smithson. Her- Quintero, M., AA. Hernandez, and S. Gaitan. 1990. Presencia del petol. Info. Sew. (54):l-70. acaro Opbionyssus natricis (Mauonyssidae) en serpientes de Moonen, J., W. Eriis, andK. van Deursen. 1979. Surinaamse slangen- un serpentarb de Mexico. Veterinaria (Mexico) 21:163-165. inkleur. C. Kersten & Co., Paramaribo. Radcliffe, C.W., D. Chiszar, and B. O'Connell. 1980. Effects of prey Moreno, E. and R Bolaiios. 1977. Hemogregarinas en serpientes de size on post-strike behavior in rattlesnakes (Cmtalus dunbus, Costa Rica. Rev. Bbl. Trop. 2547-57. C. enp, and C. oiridis). Bull. Psychon. Soc. 16:449450. Muir, J.H. 1984. Captive reproduction of the Uracoan rattlesnake, Rego, Ah. 1983a. PentastomIdeosde repteis do Brasil: revis50 dos Cmtalus uegmndis. Bull. Chicago Herpetol. Soc. 19:131-133. Cephalobaenidae. Mem. Inst. Oswaldo Cruz 78:399-411. Miiller, P. 1968. Die Herpetofauna der Insel von SoSebastlo (Bra- -1983b [1980/19811. Sobre a identificagso das esp€cies de Pom- silien). Verlag Saarbriicker Z., Saarbriicken, Germany. cepbalus (Pentastomida) que ocorrem em ofidios da Am6rica -1973. The dispersal centres of terrestrial vertebrates in the Neo- Tropical. Mem. Inst. Butantan 44/45:219-231. tropical realm. A study in the evolution of the Neotropical biota -198% [1980/19811. Notas sobre alguns pentastomldeosde rep- and itsnativelandscapes.Dr. W. Junk B.V., BiogeographicaVol. teis. Mem. Inst. Butantan 44/45233-238. 2:vi + 244 p. Renault, L. and G. Schreiber. 1951. Consideragt~esdbre albinism0 Murphy, J.B. and B.L. Armstrong. 1978. Maintenance of rattlesnakes em cascavel. Folia Clin. Biol. (So Paulo) 1691-92. in captivity. Univ. Kansas Mus. Nat. Hist. Spec. Publ. (3):vii + 40 Renjifo, J.M. 1979. Systematics and distribution of crotalid snakes in P. Colombia. MA. Thesis, Univ. Kansas, Lawrence. Nascimento, F.P. do., T.C.S. de Avila Pires, and 0.R da Cunha. 1988. Reuss, T. 1930. fjber eine neurotoxische Otterngruppe Europas, Repteis squamata de Rondania e Mato Grosso coletados atraves Mesocomnis 1927, und iiber ihre Stellung unter den Soleno- do programa Polonoroeste. Bol. Mus. Paraense Emflio Goeldi, glyphen der Welt. Glasnik Zemaljskog Muzeja, Bosni I Herce- &r. 4:21-66. govini. 4257-114. Obst, F.J., K. Richter, and U. Jacob. 1988. The completely illustrated Richardson, M. 1972. The fascinationof reptiles. Hill and Wang, New atlas of reptiles and amphibians for the terrarium. T.F.H. Publ., York. Neptune, New Jersey. River-Blanco, C. and J.R Dixon. 1979. Origin and distribution of the Oliveira, C.M. de, RP. Cardoso Junior, C.S. Brunckhorst, and A.YA. herpetofauna of the dry lowland regions of northern South Hoge. 1988. Cesariana e enterotomia em cascavel, Cmtalusdu- America, p. 281-298. In W.E. Duellman (ed.),The South Ameri- rissus collilinaatus w~peri&e- Crotalinae). Mem. Inst. Butan- can herpetofauna: its origin, evolution, and dispersal. Mus. Nat. tan 50:71-75. Hist., Univ. Kansas Monogr. No. 7. Olmo, E. 1984. Genomic composition of reptiles: evolutionary per- Roberts, M.F. 1984. Snakes. T.F.H. Publ., Inc., Neptune, NewJersey. spectives. J. Herpetol. 18:20-32. Rodrigues, E.M.B. and F.L. De Lucca. 1973. Distribution of 5SRNA in -. 1986. Animal cpogenetics. Vol. 4: Chordata 3A. Reptilia. Ge- the liver of the South American rattlesnake, Cmtalus durissur briider Bomtraeger, Berlin. temj?cus. Experientia 2957-39. Olney, P.J.S., P. Ellis, and B. Sommerfelt. 1988. Studbooks and world -and -. 1974. Ribosomal proteins from the liver of the South registers for rare species of wild animals in captivity. Int. Zoo American rattlesnake, Cmtalus durisnrs &mj?cus.Experientia Yearb. 27:482-490. 30:1407-1409. Orejas-Miranda, B. 1969. In MA. Klappenbach and B. Orejas-Miran- Roge, J.-P. and J. Sauvanet. 1987. Les serpents. Saga, Cayenne. da. Anfibios y reptiles. Nuestra Tierra (11):l-68. Roze, Jh. 1966. La taxonomia y zoogeografia de 10s ofidios en Vene- O'Shea, M. 1990. The herpetofauna of Ilha de Marad, State of Rorai- zuela. Univ. Central Venezuela, Caracas. ma, northern Brazil, p. 51-72. In J. Coote (ed.), Proceedings of -1970. Ciencia y fantasia sobre las serpientes de Venezuela. Ed. the 1988 U. K. herpetological societies symposium on captive Fondo de Cultura Cientifica, Caracas. n breeding. Brit. Herpetol. Soc., London. Ruiz, J.M. 1952[1950/19511. Sobre.a distinglo genericadosCrotalidae Pefaur, J.E. 1992. Checklist and bibliography (1960-85) of the Vene- (O~hidia:Crotaloidea) baseada emalrmnscaracteres osteol6ai-- zuelan herpetofauna. Smithson. Herpetol. Inform. Serv. (89):l- coi. Mem. Inst. Butantan 23:109-114.- 54. Sage, RD. and E.E. Capredoni. 1971. La distribuci6n de la cascabel Perez-Higareda, G. and H.M. Smith. 1991. Ofidiofauna de Veracruz; (Cmtalusdunbus temicus burentius) en Argentina y su sig- nificado zoogeo@fico (Reptilia, Serpentes). Neotropica 17: Sumichrast, F. 1881. Contribuci6n 5 la historia natural de Mhiico. I. 133-136. Notas acerca de una coleccidn de reptiles y batraciosde la pane Sanchez-Herrera,O., H.M. Smith, and D. Chiszar. 1981.Anothersug- occidental del Istmo de Tehuantepec. La Naturaleza 5268-270. gested case of ophidian deceptive mimicry. Trans. Kansas Acad. Tashjian, J. 1991. [Photograph]. The Vivarium 3(3):24. Sci. 84121-127. Taylor, E.H. 1950. Second contribution to the herpetology of San Luis Sandner Mantilla, F. 1975. Manual de las serpientes ponzoiiosas de Potosi. Univ. Kansas Sci. Bull. 35441-457. Venezuela. Ministerio de Educacih, Caracas. -1951. A brief review of the snakes of Costa Rica. Univ. Kansas -1980. Una nueva especie del genero Cmtalus (Serpentes, Cro- Sci. Bull. 343-188. talidae, Crotalinae) del sur del estado Gdrico, Venezuela. Taylor, RT., A. Flores, G. Flores, and R. Bolaiios. 1974.Geographical Mem. Cientificas Ofidiologia (5):l-12. distribution of Viperidae, Elapidae and Hydrophidae in Costa -1983 [19821. SerpientesCrotalinaedeVenezuela.Mem. Inst. Bu- Rica. Rev. Biol. Trop. 21:383397. tantan 46:193-194. Thireau, M. 1991. Types and historically important specimens of rat- Santa Rosa, CA., S. Hyakutake, P. de Biasi. H.E. Belluomini, M. Ka- tlesnakes in the Mudum National &Histoire Naturelle (Paris). warabayski, and A. Godano. 1980. ContribigBo ao estudo epi- Smithson. Herpetol. Inform. Ser. (87):l-10. demiol6gico das leptospiroses em serpents do Brasil 11. Levan- Tinoco, RA. 1978. Las serpientes de Colombia. Ciencia, mitos y ley- tamento sorol6gico em Cmtalus du&ws tanjkus Laurent, endas. Edic. Ed. Mejoras, Barranquilla. 1768 (Viperidae: Crotalinae), Cascavel. Rev. Inst. Adolfo Lutz Treviiio, C.H. 1978. Estudio herpetofaunistico distributional del sur 409-13. de Nuevo Ldn, Mexico. Biol. Thesis. Univ. Aut6n. Nuevo Mn, Savage, J.M. and J. Villa. 1986. Introduction to the herpetofauna of Monterrey. Costa Rica. Introducci6n a la herpetofauna de Costa Rica. SSAR -1980. Lista herpetol6gica anotada del sur de Nuevo Le6n, Mex., Contr. Herpetol. (3):viii + 207 p. p. 298-309. In Mem. I1 Congreso Nacional de Zoologia. Vol. 1. Schmidt, D. 1990. Schlangen. J. Neumann-Neudamm, Melsungen. Fac. Cienc. Biol., Univ. Aut6n. Nuevo Le6n, Monterrey. Schmidt, K.P. and R.F. Inger. 1957. Living reptiles of the world. Dou- Trutnau, L. 1982. Schlangen im Terrarium. Haltung, Pflege und Zucht bleday & Co., Garden City, New York. in Zwei Enden. Bd. 2. Giftschlangen. Verl. Eugen Ulmer, Stutt- -and W.F. Walker, Jr. 1943. Peruvian snakes fromthe University gan. of Arequipa. Zool. Ser., Field Mus. Nat. Hist. 24:279-296. Tryon, B.W. 1986. The island, the rattlesnake, and the Species Sur- Schnitzler, R. 1987. Die herpetologische Abteilung des Instituto Bu- vival Plan. Brit. Herpetol. Soc. Bull. 16:20-24. tantan und einige don gehaltene Schlangen. Teil3. Sauria 9:25- Tu,A.T. 1971. The mechanism of snake venom actions-rattlesnakes 30. and other crotalids, p. 87-109. In L.L. Simpson (ed.), Neuropoi- Schreiber, G., N.B. Melucci, S.E. Gerken, Y.X Sant'Ana, LA. Fallieri, sons: their pathophysiological actions. Vol. 1. Poisonsof animal and F.O. Amorim. 1967 [19661. Pequisas de citologia quantita- origin. Plenum Press, New York. tiva.WC - DNAe volume nuclear nos tecidossom6ticosdosver- -. (ed.). 1982. Rattlesnake venoms. Their actions and treatment. tebrados. Mem. Inst. Butantan 33135-150. Marcel Dekker, Inc., New York. Scott, N.J. and J.W. Lovett. 1975. A collection of reptiles and amphibi- Vanzolini, P.E., A.M.M. Ramos-Costa, and L.J. Via. 1980. Repteis das ans from the Chaco of Paraguay. Univ. Connecticut Occ. Pap., caatingas. Acad. Brasileira Cienc., Rio de Janeiro. Biol. Sci. Ser. 2:257-266. Vellard, J. 1946. Morfologiadel hemipenis yevolucidn delosofidios. Scudder, K.M., D. Chiszar, and H.M. Smith. 1983. Effect of environ- Acta Zool. Lilloana 5263288. mental odors on strike-inducedchemosensorysearchingby rat- Villa, J. 1984. The venomous snakes of Nicaragua: a synopsis. Mil- tlesnakes. Copeia 1983:519-522. waukee Pub. Mus. Contr. Biol. Geol. (59):l-41. Seba, A. 1735. ~ocu-pletissimirerum naturalium thesauri accurata de- -, L.D. Wilson, and J.D. Johnson. 1988. Middle American herpe- scriptio et iconobis artificiosissimis expressio, per universam tology. A bibliographic checklist. Univ. Missouri Press, Colum- phisices historiam. Vol. 2. J. ~estenium;~. mi&, andJanson- bia. io-Waesbergios, Amsterdam. Vitt, L.J. and L.D. Vangilder. 1983. Ecology of a snake community in Smith, A.W., M.P. Anderson, D.E. Skilling, J.E. Barlough, and P.K. northeastern Brasil. Amph.-Rep. 42732%. Ernsley. 1986. First isolation of calcivirus from reptiles and am- Wagler, J. 1824. Serpentum Brasiliensium species novae ou histoire phibians. Amer. J. Vet. Res. 47:1718-1721. naturelle des espi?cesnouvellesde serpens, recueillies et obser- Smith, H.M. and G. Perez-Higareda. 1987. The literature on somato- v&s pendant le voyage dans I'interieur du Br&il dans les anne- dichotomy in snakes. Bull. MarylandHerpetol. Soc. 23139-153. es 1817,1818,1819,1820, execute par ordre de Sa Majestt le Roi -and RB. Smith. 1976. Synopsis of the herpetofauna of Mexico. de BavKre, publiee par Jean de Spix, &rite d'aprh les notes du Vol. 111. Source analysis and index for Mexican reptiles. John voyageur. Franc. Seraph. Hiibschmanni, Monachii. Johnson, North Bennington, Vermont. Watt, G. 1989. Snakebitetreatment and first aid, p. 6-13. In JA. Camp- -and -. 1993. Synopsis of the herpetofauna of Mexico. Vol. bell and W.W. Lamar. The venomous reptiles of Latin America. VII. Bibliographic addendum IV and index, bibliographic ad- Comstock Publ. Assoc., Ithaca, New York. denda II-IV 1979-1991. Univ. Press Colorado, Niwot. Wellborn, S., K.M. Scudder, H.M. Smith, K. Stimac, and D. Chiszar. -and E.H. Taylor. 1945. An annotated checklist and key to the 1982. Investigatory behavior in snakes 111: effects of familiar snakes of Mexico. Bull. U.S. Natl. Mus. (18i?:iv + 239 p. odors on investigation of clean cages. Psychol. Rec. 32169-177. -and-. 1950. Type localities of Mexican reptiles andamphibi- Werman, S.D. 1992. Phylogenetic relationships of Central and South ans. Univ. Kansas Sci. Bull. 35313-380. American pitvipers ofthe genus ~otbm&(sew lato):cladistic Snider,A.T. andJ.K. Bowler. 1992. Longevityof reptiles andamphibi- analvses of biochemical and anatomical characters. D. 21-40. In ans in North American collections. 2nd ed. SSAR Herpetol. Circ. ~A.dam~belland E.D. Brodie, Jr. (eds.), Biology ;df the pitvi- (21):iii + 40 p. pers. Selva, Tyler, Texas. Sol6nan0,A. and L. Cerdas. 1988. Biologia reproductiva de la casca- Wilson, L.D. and J.R. McCranie. 1991. Additional departmental re- be1 centroarnericana Cmtalus durissuc durissuc (Serpentes: Vi- cords for the herpetofauna of Honduras. Herpetol. Rev. 2269- peridae) en Costa Rica. Rev. Biol. Trop. 36:221-226. 71. Sonnini, C.S. and PA. Latreille. 1801. Histoire naturelle des reptiles, -and J.R. Meyer. 1985. The snakes of Honduras. 2nd ed. Milwau- avec figures dessinees d'aprss nature. Seconde partie. Serpens. kee Pub. Mus., Milwaukee, Wisconsin. Vol. 3. Deterville, Paris. Yarrow, H.C. 1875. Report upon the collections of batrachians and StanEk, V.J. 1962. Introducing poisonous snakes. Golden Pleasure reptiles made in portionsof Nevada, Utah, California,Colorado, Books, London. New Mexico, and Arizona, during the years 1871,1872,1873, Staton, MA. andJ.R. Dixon. 1977.The herpetofauna ofthe central Lla- and 1874. Rept. Geog. Geol. Exped. Surv. W. 100 Meridian, nos of Venezuela: noteworthy records, a tentative checklist and Wheeler 5509-584. ecological notes. J. Herpetol. 11:17-24. Stille, B. 1987. Dorsal scale microdermatoglyphics and rattlesnake James R McCnnie, 10770 SW 164 Street, Miami, FL 33157, USA. (Cmtalus and Sistmnrs) phylogeny (Reptilia: Viperidae: Crotal- inae). Herpetologica 4598-104. Primary editor for this account, Larry David Wilson. Sullivan, B. 1974. Reptilian hemoglobins, p. 377-398. In M. Florkin and B.T. Scheer (eds.), Chemical zoology. Volume IX Arnphib- Published 30 November 1993andcopwight O 1993bythe Society for ia and Reptilia. Academic Press, New York. the Study of Amphibians and ~e~tilds..-