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Bothrops atrox (LINNAEUS, 1758)

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Higher TaxaViperidae, Crotalinae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Common NamesE: Barba Amarilla, Fer-de-Lance, common lancehead
G: Caicaca, Gewöhnliche Lanzenotter
NL: Gewone lanspuntslang
Portuguese: Acuambóia, Boca-Podre, Cambéua, Caiçaca, Caiçara, Comboia, Cuamboia, Japoboia, Jaracuçu, Jararaca, Jararaca-Açu, Jararaca-da-Amazônia, Jararaca-do-Amazonas, Jararaca-do-Norte, Jararaca-do-Rabo-Branco, Jararaca-Grão-de-Arroz, Jararacarana, Mapanare, Surucucu, Surucucu-da-Várzea, Surucucu-do-Barranco, Surucucurana
S: Mapanare; Macagua; Guayacán; Talla X; Cuatro narices; Rabo amarillo 
SynonymColuber atrox LINNAEUS 1758: 222
Coluber lanceolatus LACÉPÈDE 1789
Bothrops furia WAGLER 1824: 52
Bothrops sabinii GRAY 1842: 47
Bothrops subscutatus GRAY 1842: 47
Trigonocephalus colombiensis HALLOWELL 1845: 246
Bothrops affinis GRAY 1849: 7
Bothrops atrox — DUMÉRIL & BIBRON 1854: 1507
Craspedocephalus atrox — WUCHERER 1863: 242
Bothrops asper GARMAN 1883
Lachesis lanceolatus — BOULENGER 1896
Bothrops lanceolatus — GARCIA 1896: 22
Trigonocephalus asper GARMAN 1883 (fide SCHÄTTI & KRAMER 1993)
Lachesis atrox — QUELCH 1899
Bothrops neuvoiedii venezuelenzi BRICEÑO 1934: 46
Trimeresurus atrox — POPE 1955
Bothrops atrox — PETERS & OREJAS-MIRANDA 1970: 44
Bothrops isabelae SANDNER MONTILLA 1979
Bothrops atrox — GASC & RODRIGUES 1980
Bothrops lanceolatus aidae SANDNER MONTILLA 1981
Bothrops lanceolatus nacaritae — SANDNER MONTILLA 1981
Bothrops lanceolatus nacaritae — SANDNER MONTILLA 1982
Bothrops atrox aidae VANZOLINI 1986
Bothrops lanceolatus nacaritae SANDNER MONTILLA 1990
Bothrops atrox — WELCH 1994: 31
Bothrops isabelae — KORNACKER 1999: 169
Bothrops atrox — MCDIARMID, CAMPBELL & TOURÉ 1999: 256
Bothrops atrox — MONZEL & WÜSTER 2008
Bothrops isabelae — MONZEL & WÜSTER 2008
Bothrops isabelae — FENWICK et al. 2009
Bothrops atrox — FENWICK et al. 2009
Bothrops atrox — CARRASCO et al. 2012
Bothrops atrox — WALLACH et al. 2014: 116
Bothrops isabelae — WALLACH et al. 2014: 117
Bothrops atrox — NATERA-MUMAW et al. 2015: 244
Bothrops colombiensis — NATERA-MUMAW et al. 2015: 246
Bothrops isabelae — NATERA-MUMAW et al. 2015: 247
Bothrops colombiensis —SEÑARIS et al. 2018: 302
Bothrops atrox — NOGUEIRA et al. 2019 
DistributionGuyana, Suriname, French Guiana, Venezuela, Brazil (Pará, Acre, Amazonas etc.), Colombia, E Ecuador, NE/E Peru, N Bolivia, Trinidad

Type locality: “Asia” (in error); true locality unknown fide PETERS 1960.

colombiensis (invalid): N Venezuela; Type locality: Republic of Colombia, within 200 miles of Caracas.

aidae (invalid): Venezuela; Type locality: “Selvas de Guatopo, Estado Miranda”

isabelae (invalid): Venezuela; Type locality: "siete kilometros al SE. de Guanare, Estado Portuguesa, a pocos metros de la carretera que va de Guanare a Guanarito '" a 182 mts. sobre el nivel del mar" [= 7 km southeast of Guanare, Estado Portuguesa, Venezuela, elevation 182 m.].

nacaritae (invalid): Venezuela; Type locality: “Cariprima, N. de Valencia"  
Reproductionovovivparous; facultative parthenogenesis (Cubides-Cubillos et al. 2020) 
TypesSyntypes: NRM (NHRM) 100-101, was "Mus. Ad. Fr.".
Syntypes (2): ANSP 6974 and ANSP 6975 [colombiensis]
Holotype: "Live specimen in the Serpentario del Instituto Venezolano de Ofidiologia, Caracas, 15-61979." [isabelae]
Holotype: ZSM, uncatalogued specimen(s) (lost) [furia] 
DiagnosisDiagnosis: With the possible exception of a few problematic species of the Bothrops atrox complex (see Remarks), B. atrox is distinguished from all congeners and from species of Bothrocophias and Bothriopsis by the following combination of characters: (1) lacunolabial present; (2) dorsal keels nontuberculate; (3) infralabials and gulars pigmented to varying degrees but lacking regular series of cream round to C-shaped marks; (4) canthorostrals absent; (5) subcaudals divided; (6) ventrals 182–200 and subcaudals 54–72 [Campbell and Lamar 1989 report 169–212 ventrals and 47–86 based on literature records from across the range of the species complex. Their wider ranges are based on published accounts that may not apply to the species occurring in Bolivia. Our definition is based on specimens we examined]; (7) loreal subtriangular, higher than long; (8) canthals narrow and separated by 3–7 scales; (9) internasals usually in contact, separated by one scale in 22% of the sample; (10) postocular stripe distinct and brown; two temporals high; (11) dorsum brown, pattern of subtriangular blotches or French telephone-shaped marks; venter uniformly and heavily speckled (populations on Andean slopes) or with alternating cream and charcoal, square blotches (Harvey 2005: 16). 

Synonymy: Synonymy partly after PETERS & OREJAS-MIRANDA 1970. Sandner-Montilla (1979a) synonymised B. colombiensis with B. lanceolatus. Not listed as such by WELCH 1994 or mentioned by FENWICK et al. 2009. Listed as synonym of B. atrox by KORNACKER 1999 and McDiarmid et al. 1999. RIVAS et al. 2012 consider the populations from Valencia and Guatopo as B. asper rather than B. atrox, except the population from the cloud forest that is B. venezuelensis. RIVAS et al. 2012 consider B. colombiensis as synonym of B. atrox. B. atrox may include a number of other nominal species, especially B. colombiensis, B. isabelae, B. leucurus, B. marajoensis, B. moojeni, B. pradoi. In addition, B. atrox has often been confused with B. asper.

Bothrops isabelae has been synonymized with B. atrox by MARKEZICH & TAPHORN 1993 and other authors but used as a valid species by Salomão et al. (1999) and others. Placed in the synonymy of Bothrops atrox atrox by Golay et aI. 1993 and McDiarmid et al. 1999. In all analyses by Fenwick et al. (2009), B. isabelae falls together with B. atrox, with a seemingly low genetic distance. Nevertheless, Fenwick et al. (2009) included B. isabelae in their list of Bothrops species (therefore apparently recognizing it as a valid taxon). RIVAS et al. 2012 consider B. isabelae as synonym of B. atrox. See also McDIARMID et al. (1999) for discussion.

Distribution: STUART (1935) reported this species from Guatemala (probably in error). MARTIN (1958) listed it for Mexico (Tamaulipas). “Cariprima” is not found on any available map, but both Estado Miranda and Valencia are in the Coastal Range [RIVAS et al. 2012]. See map in Harvey 2005: 14 (Fig. 9). See map in Nogueira et al. 2019.

Illustrations: Penner (2001). Plate 111 (p. 414) in Stebbins & McGinnis (2012) shows a Crotalus scutulatus, not C. atrox. 
EtymologyNamed after Latin atrox = hideous, savage. 
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