Bothrops, WAGLER, 1824

Fenwick, Allyson M., Gutberlet Jr, Ronald L., Evans, Jennafer A. & Parkinson, Christopher L., 2009, Morphological and molecular evidence for phylogeny and classification of South American pitvipers, genera Bothrops, Bothriopsis, and Bothrocophias (Serpentes: Viperidae), Zoological Journal of the Linnean Society 156 (3), pp. 617-640 : 631-632

BOTHROPS WAGLER, 1824

Type species: Bothrops lanceolatus (Bonnaterre, 1790)

Etymology: The generic name is derived from the Greek bothros, referring to the facial pit, and ops, meaning either ‘eye’ or ‘face’. It refers to the loreal pit between the nostril and eye, and names ending in this suffix are masculine.

Content: Bothrops andianus , B. asper , B. atrox , B. brazili , B. caribbaeus , B. isabelae , B. jararacussu , B. lanceolatus , B. leucurus , B. marajoensis , B. moojeni , B. muriciencis , B. osbornei , B. pirajai , B. punctatus , B. sanctaecrucis , and B. venezuelensis .

Definition: Members are of moderate length to elongate, are thin to moderately stout, and are terrestrial, lacking a prehensile tail. Dorsal colour brown to black, with trapezoidal to spade-shaped markings on most species ( B. lanceolatus with spots, B. osbornei and B. punctatus with vertical bands). Head pattern variable, from patternless, to speckled, to paired spots, to spadeshaped pattern, showing a postorbital stripe in most species (faint to absent in B. brazili and B. sanctaecrucis , absent in some B. moojeni ); there are no other distinctive head markings.

There are 3 or 4 interoculabials, 7 or 8 supralabials, 3–13 smooth or keeled intersupraoculars, 3–11 scales between the first pair of postcanthals, 24–36 interrictals, 153–227 ventrals, 22–33 dorsal scale rows at midbody, and 38–72 divided subcaudals (one B. atrox and two B. jararacussu specimens with both divided and entire subcaudals). Prelacunal and second supralabial fused (one B. brazili specimen with scales divided), supralacunal separate from middle preocular (one B. asper and one B. atrox with scales fused). Sublacunal entire, loreal pit ventral to naso-orbital line (one B. caribbaeus and one B. venezuelensis with pit crossed by line). Dorsal scales keeled with typical thin ridge.

From an examination of the hemipenes of B. atrox , B. asper , B. brazili , B. jararacussu , B. leucurus , B. moojeni , B. punctatus , and B. venezuelensis : many lateral spines, lateral calyces distal to crotch (one quarter of B. brazili specimens with lateral calyces reaching crotch).

From an examination of osteological samples of B. asper , B. atrox , B. brazili , B. jararacussu , B. moojeni , and B. punctatus : pleurapophyses of midcaudal vertebrae long and slender (one-quarter of B. brazili specimens with short and slender pleurapophyses), 3–5 palatine teeth, 12–21 pterygoid teeth, and 8–18 dentary teeth. Maxillary fang longer than height of maxilla, well-developed medial wall of pit cavity in maxilla, pit in anterolateral wall of maxillary pit cavity simple or with a small rounded projection. Lateral margin of head of ectopterygoid intermediate to narrow, shaft of ectopterygoid flat and tapering or narrow without tapering, pits on posterior surface of anterior end of ectopterygoid single or paired, ectopterygoid base long and overlapping, base of ectopterygoid longer than base of pterygoid. Choanal process of palatine positioned medially, moderate to attenuate in shape. Medial margin of dorsal portion of prefrontal moderately to weakly concave, dorsal surface of frontals with elevated margins (one specimen of B. asper and one of B. atrox with flat dorsal surface). Supratemporal with a small projection (one B. asper with expanded supratemporals lacking projections); supratemporal thick and rounded. Single Meckellian foramen.

Diagnosis: Bothrops differs from other South American pitvipers in 50 mitichondrial characters ( Table 4). In addition, Bothrops species generally have four palatine teeth, which is a morphological synapomorphy of the genus ( B. moojeni and B. jararacussu have five; B. brazili and B. sanctaecrucis have three). Bothriopsis and Bothrops are distinguished from other KEY TO SOUTH AMERICAN BOTHROPOID GENERA

1. Dorsal ground colour green, grey, or brown, dorsal head colour black or matching dorsum, tail prehensile, prelacunal and second supralabial fused................................................................................................2 Dorsal ground colour and dorsal head colour gold or brown to black, tail not prehensile, prelacunal and second supralabial fused or separate, with no or one row of subfoveals.................................................................3

2. Found east of the Andes, dorsal colour usually green (lavender grey to green in Bothriopsis taeniata , tan, brown, grey, or olive in Bothriopsis medusa ) ....................................................................................... Bothriopsis Found west of the Andes, dorsal colour brown to greenish tan ........................................................ Bothrops

3. Keel on dorsal scales tuberculate on caudal part of body, rostral higher than broad or square, distinct white spots on posterior infralabials and gulars may be present, canthorostrals may be present, 125–169 ventral scales (one specimen with 192 scales),................................................................................................. Bothrocophias Keel on dorsal scales typical thin ridge, rostral broader than high to square, or higher than broad in species lacking tuberculate dorsal scales, distinct white spots and canthorostrals absent, 145–227 ventral scales....................4

4. Black bars on gular scales may be present; if absent, species has dorsal pattern of spots or parallel bands, and nonprehensile tail. Dark patterning on head generally spade-shaped; head has a pattern of paired spots in species that have black bars on gular scales, have a dorsal pattern of parallel bands, or lack a nasal pore. Prelacunal and second supralabial separate with no or one subfoveal, loreal scale longer than high to higher than long, 25–40 interrictals, 25–55 subcaudals............................................................................................. Rhinocerophis Black bars on gular scales absent, markings on dorsum trapezoidal to spade shaped, except in species with prehensile tails. Dark patterning on head absent, speckled, as paired spots, or spade shaped. Prelacunal and second supralabial fused or separate, with either no or one subfoveal, loreal scale longer than high to square, 21–34 interrictals, 31–72 subcaudals..............................................................................................................5

5. Prelacunal and second supralabial separate, with no or one subfoveal scale; if fused, species is a Brazilian island endemic ( B. alcatraz or B. insularis ) or a coastal mainland species in southern Brazil, northeastern Paraguay, and northern Argentina, generally having 8 supralabials and 170–216 ventrals ( B. jararaca ). Subcaudals both divided and entire, or all divided, 7–11 supralabial scales, 144–206 ventral scales, mesial spines on hemipenes present ...................................................................................................................................... Bothropoides Prelacunal and second supralabial fused. Species sympatric with B. jararaca either have fewer supralabials or fewer ventrals, or both. In all species, subcaudal scales divided, 7 or 8 supralabial scales, 153–227 ventral scales, mesial spines on hemipenes absent or present.............................................................................. Bothrops

South American genera by having higher numbers of ventrals (157–236 and 153–227, respectively, compared with 125–206), and by having the prelacunal fused to the second supralabial (also seen in Bothropoides jararaca , B. alcatraz , B. insularis , and in some Bothrocophias ). Bothrops is distinguished from Bothriopsis in its brown to black coloration and lack of a prehensile tail, except for Bothrops osbornei and Bothrops punctatus with prehensile tails. These two Bothrops species occur west of the Andes, as opposed to Bothriopsis species that all range east of the Andes.

Distribution: Most species are found in South America east of the Andes, exclusive of Uruguay, southern Paraguay, and central to southern Argentina ( Campbell & Lamar, 2004). Bothrops caribbaeus and B. lanceolatus are found on the Caribbean islands of Saint Lucia and Martinique. Bothrops osbornei , B. punctatus , and B. asper range through Peru, Ecuador, and portions of Colombia west of the Andes, and B. asper ranges northwards in Middle America through the countries of Panama, Costa Rica, Nicaragua, Honduras, Guatemala, Belize, and Mexico. See Campbell & Lamar (2004) for range maps of individual species. Remarks: According to Ferrarezzi & Freire (2001), in Campbell & Lamar (2004), Bothrops muriciencis is most similar in overall appearance to Bothrops pirajai , B. brazili , B. jararacussu , and B. sanctaecrucis , with B. pirajai suggested as the closest relative. Bothrops pirajai is poorly known, and specimens were unavailable, but it is very similar to some specimens of B. brazili and B. jararacussu ( Campbell & Lamar, 2004) . As the aforementioned species included in the study all are found in Bothrops , as described in this paper, we assign B. muriciencis and B. pirajai to the genus as well.