Bothropoides, Fenwick & Gutberlet Jr & Evans & Parkinson, 2009

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 : 629-630

BOTHROPOIDES GEN. NOV.

Type species: Bothrops neuwiedi Wagler, 1824 .

Etymology: The generic name is derived from the Greek bothros, referring to the facial pit, and also referring to the currently named genus Bothrops . The term oides means ‘similar to’ or ‘having the nature of ’, thereby recognizing the affinity of these species with other terrestrial South American pitvipers. Names ending in this suffix are masculine.

Content: Bothropoides alcatraz , B. diporus , B. erythromelas , B. insularis , B. jararaca , B. lutzi , B. mattogrossensis , B. neuwiedi , B. pauloensis , and B. pubescens .

Definition: Members are of moderate length and girth, and are terrestrial, lacking a prehensile tail. Dorsal colour gold ( B. insularis ) to brown or black, with spade-shaped dorsal markings, with some lacking spots between the spades ( B. alcatraz , B. insularis , B. jararaca , B. pauloensis , and B. diporus ), and with others showing them ( B. erythromelas , B. lutzi , B. mattogrossensis , B. neuwiedi , and B. pubescens ). A postorbital stripe is present (but is pale in most B. insularis specimens); dorsal head patterning is variable among species, and they share no other distinctive head markings.

There are 3–5 interoculabials, 7–11 supralabials, 5–12 keeled intersupraoculars (smooth in B. erythromelas and one specimen each of B. insularis and B. alcatraz ), 4–10 scales between the first pair of postcanthals, 21–34 interrictals, 144–206 ventrals, 21–30 dorsal scale rows at midbody, and 31–66 divided or divided and entire subcaudals. The prelacunal and second supralabial are fused (in B. jararaca , B. alcatraz , and B. insularis ) or separate, with 0–1 rows of subfoveals. Supralacunal separate from middle preocular (one B. mattogrossensis had scales fused). Loreal wider than high or square (one B. neuwiedi had loreal higher than wide), loreal pit ventral to nasoorbital line. Postnasal in contact with first supralabial in some individuals. Dorsal scales keeled with typical thin ridge.

From an examination of hemipenes of B. diporus , B. alcatraz , and B. insularis : many lateral spines on hemipenes with lateral calyces distal to crotch in most members of the genus, and few lateral spines with lateral calyces reaching crotch in B. insularis . Mesial spines present on hemipenes, except for half of the B. insularis specimens. Calyces spinulate, except in one B. insularis with smooth calyces.

From an examination of osteological samples of B. neuwiedi and B. jararaca : 3–5 palatine teeth, 10–16 pterygoid teeth, and 11–15 dentary teeth. Maxillary fang longer than height of maxilla, well-developed medial wall of maxillary pit cavity, with pit in anterolateral wall of maxillary pit cavity either simple or with a small rounded projection. Foramen absent from ventral surface of lateral process of prootic. Lateral margin of head of ectopterygoid of intermediate width, ectopterygoid shaft flat and tapering or narrow without tapering, and ectopterygoid base with a long overlapping projection. Choanal process of palatine positioned medially, and greatly reduced ( B. neuwiedi ) or attenuate ( B. jararaca ) in shape. Meckellian foramen single; angular and splenial partially fused.

Diagnosis: Bothropoides differ from other South American pitvipers in 38 mitochondrial characters ( Table 4). External characters overlap with other South American genera, with no unique synapomorphies in scalation. Distribution in eastern South America, combined with terrestrial habitat in grasslands or broad-leaf forests ( Bothropoides neuwiedi group), or semiarboreal habitat in Atlantic forests ( B. jararaca group), distinguishes this genus from others (see Table 4). Bothropoides has fewer interrictals (21– 34) than the other South American genera (24–40), and some individuals have high numbers of supralabials (7–11, also seen in Rhinocerophis ; all other South American genera have 7 or 8 supralabials). Bothropoides differs from Bothrops and Bothriopsis in having most species with the prelacunal separate from the second supralabial ( B. jararaca , B. alcatraz , and B. insularis have the prelacunal fused to the second supralabial). Some specimens have both divided and entire subcaudals, a state also seen in Bothriopsis . Bothropoides differs from Bothriopsis in the lack of a prehensile tail and lack of green coloration. It differs from Bothrocophias in the lack of white spots on the gular scales, and the lack of tuberculate keels on posterior dorsal scales. Bothropoides differs from some Rhinocerophis ( R. alternatus , R. cotiara , R. fonsecai , and R. jonathani ) in the absence of distinctive back bars on the underside of the head.

Distribution: Eastern South America: in Brazil and associated islands, Bolivia, south-eastern Peru, Paraguay, Uruguay, and northern to central Argentina ( Campbell & Lamar, 2004). See Campbell & Lamar (2004) for range maps of individual species.

Remarks: We did not examine individuals of Bothrops lutzi , but based on prior work that elevated this species out of the Bothrops neuwiedi complex ( Silva, 2000, 2004), we include it in the genus Bothropoides .