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Dipsas bothropoides MEBERT, PASSOS, FERNANDES, ENTIAUSPE-NETO, QUEIROZ-ALVEZ, MACHADO & LOPES, 2020

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Higher TaxaColubridae (Dipsadinae), Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Subspecies 
Common Names 
SynonymDipsas bothropoides MEBERT, PASSOS, FERNANDES, ENTIAUSPE-NETO, QUEIROZ-ALVEZ, MACHADO & LOPES 2020 
DistributionBrazil (Minas Gerais, probably Bahia)

Type locality: “Fazenda Duas Barras” (near 16°25′0.00′′S, 40°2′60.00′′W, 800 m elevation), a ranch in the district of Talimã, municipality of Santa Maria do Salto, Minas Gerais, Brazil  
Reproduction 
TypesHolotype: MNRJ 26377, adult female, collected on 8 November 2004 by Daniel S. Fernandes and Luciana B. Nascimento.
Paratype: MZUESC 15828, an adult male found freshly killed on state road BA-001 on 23 January 2016 by Konrad Mebert and Carol Cornélio, 2 km northeast of Cachoeira do Tijuipe and 3 km southwest of Txai, Praia (Beach) de Itacarézinho (14°23′43.14′′S, 39°2′21.70′′W; 53 m a.s.l.), Itacaré, Bahia, Brazil (Figs. 3, 4). We collected no tissue sample from the holotype, but such is available from the paratype at MZUESC. 
DiagnosisDiagnosis: The new species is allocated to the genus Dipsas by possessing the following derived characters combined from Peters (1960), Cadle (2007), and Arteaga et al. (2018): mental sulcus (groove) absent, shape of chinshields (square or polygonal rather than elongate and narrow), more than two pairs of chinshields, absence of supralabial noticeably higher than other supralabials and in contact with postocular, and Harderian gland occupying the entire postorbital region.
Dipsas bothropoides sp. nov. (holotype and paratype) is distinguished from all currently recognized congeners by the unique combination of the following morphological characters: (1) dorsal scale rows 15/15/15, smooth; (2) one pair of infralabials in contact posterior to mental; (3) infralabials 8–9, 4–6th or only 4th in contact with second pair of chinshields; gulars separating infralabials from preventrals and ventrals; (4) supralabials 8–9 (4–5th or 5–6th contacting orbit); (5) nasal partially or fully divided; (6) internasal paired or fused; (7) loreal square usually in contact with orbit, but contact can be obstructed by a tiny interjacent scale; (8) preoculars 1 or 2 present above and/or below loreal, excluding prefrontal from orbit; (9) postoculars two, excluding temporals from orbit; (10) temporals 3/3/3, 2/3/2 or 2/3/3; (11) ventrals 183 in female (holotype), 179 in male (paratype); (12) subcaudals 100 in female (holotype) and 110 in male (paratype); (13) head light brown with 3–7 large, black blotches with light to yellow borders on the pileus, including a preorbital band (holotype) or three blotches instead (paratype), with the paratype exhibiting two additional parallel spots diverging anteriorly along the sutures of parietals-frontalsupraoculars; (14) first dorsal blotch (nuchal collar) 2–5 scales long on the mid-dorsals and 10–14 scales long at paraventral portion; (15) dorsum of body medium brown (paratype before preservation) to light brown (preserved holotype) with well-defined dorsal blotches in the shape of triangles with black (paratype before preservation) to dark brown borders, becoming lighter towards the center of the blotch, and an additional cream to yellow border on the flanks (both specimens) and venter (paratype), more prominent anteriorly in the male before preservation; (16) number of dorsal blotches 20–26 ( = 23; SD = 2.9; n = 4 sides) to vent, arranged in pairs and separated by a light, slightly enlarged row of vertebral scales; (17) midlevel of dorsal blotches 4–7 scales long; (18) number of dorsal blotches on the tail 10–16 ( = 13; SD = 2.9; n = 4 sides), dark brown with yellow-cream border, fading increasingly in posterior blotches; (19) venter cream brown anteriorly, becoming variably darker through stippling towards posterior region of body and tail up to exhibit the same coloration as dorsum; (20) maxillary teeth 16–17; (21) pterygoid teeth 16–17; (22) palatine teeth 9/9; (23) hemipenis unilobed with conspicuous capitular arch on asulcate side; (24) capitulum on asulcate face of hemipenis with two lobular longitudinal crests, oriented obliquely and converging medially to tip of organ.
The general body habitus of Dipsas bothropoides sp. nov. and its elongate head shape, in particular the preorbital-snout area, as well as some features of color pattern resemble members of the D. incerta species group (sensu Harvey, 2008; Fernandes et al., 2010). This group currently includes D. sazimai, D. alternans, D. incerta Jan, 1863 (holotype depicted in Jan and Sordelli [1870], reprinted in Passos et al. [2004]), and D. praeornata Werner, 1909. The D. incerta group is further defined by having a brown dorsal ground color with darker blotches, a mostly immaculate head with few blotches, 15 dorsal scale rows, and loreal in contact with the orbit, all of which are consistent with our specimens. Although Harvey (2008) and Fernandes et al. (2010) stated that the nuchal collar does not reach the rictus in members of the D. incerta group, a D. alternans specimen depicted in Maia-Carneiro et al. (2012) appears to show a nuchal collar that may reach the rictus, as it does in the new species.

Comparison of external characters within the Dipsas incerta group: Dipsas bothropoides sp. nov. differs from D. incerta (Passos et al., 2004: fig. 4) and D. praeornata (Fig. 7D) by its triangular dorsal blotches, dark iris, and 9 infralabials (vs. rhomboid to ovoid dorsal blotches, light iris, and 11–14 infralabials). Furthermore, D. incerta and D. praeornata have been documented only from several sites within Venezuela and the Guiana region, 2,000–3,000 km from the new species (Jan, 1863; Harvey, 2008; Lotzkat et al., 2008; Natera-Mumaw et al., 2015), and thus can be safely excluded as being conspecific with D. bothropoides sp. nov. In the latter two publications, D. praeornata is listed and treated as D. latifrontalis. However, D. latifrontalis belongs to the Dipsas peruana group (Arteaga et al., 2018), and until a more detailed analysis clarifies the problem, we assign all D. latifrontalis from those two publications to D. praeornata, except for the holotype, BMNH 1946.1.20.98.
Dipsas bothropoides sp. nov. resembles D. alternans in all scalation characters (see Table 1, Figs. 7B, 8D, Passos et al. [2004], and details below), but differs from it in the following parameters: color pattern, including black dorsal triangular blotches with light centers and yellow-cream borders fading towards the tail tip (vs. rounded dorsal dark brown blotches that may have a white border); 3–7 large, yellow-bordered pileus blotches on top of head (vs. 2–3 large, yellow-bordered blotches, although some specimens display several small black spots with no light border); groups of tightly arranged square brown ventral spots, increasing posteriorly along lateral margins of ventrals, and alternating with groups of spots closer to mid-venter (vs. fewer and more loosely arranged groups of ventral spots, often triangular and narrower, predominantly along lateral margins of ventrals). Prefrontals not in contact with orbit, as in D. alternans, although contact occurs occasionally in the latter species (e.g., D. alternans IBSP 22889). However, D. alternans is known only from southeastern Brazil, with the nearest records ca. 360 km south at Baixo Guandu (IBSP 9280) and 400 km at Santa Teresa (MNB R704), Espírito Santo (Fig. 3; Peters, 1960; Passos et al., 2004).
From the Dipsas incerta group, only D. sazimai is known to occur within or near Brazil’s northeastern Atlantic coastal belt (Fig. 3) and might be sympatric with D. bothropoides sp. nov. in mountains up to 100 km from the Atlantic coast of northeastern Brazil (Fernandes et al., 2010; Roberto et al., 2014), but not at the lowland locality of the new species. Dipsas bothropoides sp. nov. differs from D. sazimai in the following characters (Figs. 2, 7A, C, 8A–C): prefrontals not contacting orbit (vs. prefrontals in contact with orbit); gulars separating second pair of chinshields from infralabials (vs. infralabials in direct contact with second pair of chinshields); triangular dark-edged dorsal blotches with light centers slightly shorter than interspaces throughout the body and tail (vs. rounded dorsal dark brown blotches generally wider than interspaces becoming lighter and narrower posteriorly, some with a thin vertical line with an increasing size of interspaces); immaculate interspaces throughout body (vs. posterior interspaces can exhibit a thin vertical line); 3–7 large pileus blotches with yellow borders (vs. variable number of medium to large black blotches without yellow border, although some specimens have several small black spots instead); groups of brown, square ventral spots along lateral margins of ventrals, alternating with similar but smaller groups closer to mid-venter (vs. groups of ventral spots, or spots forming an uninterrupted, continuous line only along lateral margins of ventrals, not alternating with mid-venter brown groups, spots increasing in size posteriorly in some specimens).

Comparison of external characters with sympatric arboreal Dipsas species
We compared D. bothropoides sp. nov. with four sympatric Dipsas species (see Argôlo, 2004): (1) D. catesbyi Sentzen, 1796; (2) D. variegata (including former D. neivai, which represents the Atlantic Forest populations of the widespread D. variegata; Harvey and Embert, 2008); (3) D. indica sensu lato; and (4) D. albifrons (Dias et al., 2018). Dipsas bothropoides sp. nov. differs from D. catesbyi (Figs. 6G, 7H) by: yellow-brown head with a few yellow-bordered black blotches (vs. strongly contrasting black-and-white color pattern on the head with a black-cap extending below the eye); dorsal ground color brown with triangular light centered blotches and a relatively constant blotch/interspace ratio posteriorly (vs. large black-and-white transversal bands anteriorly changing posteriorly to a red-brown body color with oval black bilateral blotches or saddles, blotch/interspace ratio decreasing posteriorly); loreal contacting the orbit, two temporals posterior to postoculars, and vertebral scales only moderately enlarged, 1.4 times larger than adjacent dorsals (vs. loreal not contacting orbit, usually single first large temporal [but see variation among D. catesbyi from the Amazon in Lima and Prudente, 2009], and large expanded vertebral scales 2–3 times wider than adjacent dorsal scales); dorsals 15 (vs. 13 in D. catesbyi); ventrals 179–183 (vs. 186–201 in Bahia specimens of D. catesbyi, in contrast to D. catesbyi males from the Amazon with ventrals as few as 160, see Lima and Prudente, 2009).
The new species differs from Dipsas indica (Figs. 7I, 8G), D. variegata (Figs. 7F, J, 8E), and D. albifrons (Figs. 7E, 8F) by: head more elongate (see Peters, 1960 for description of head truncation in these species), particularly preorbital area (vs. a comparatively shorter, but higher snout, and larger eyes resulting in a convex supraocular) and with corresponding different cephalic scale arrangement. For example, D. bothropoides sp. nov. exhibits predominantly square labials, and only two first infralabials narrow and vertically elongate (vs. anterior 2–5 supralabials and 2–6 infralabials are narrow and tall in D. indica, D. varieagta, and D. albifrons); first pair of infralabials contacting each other behind mental scale (vs. 1st and 2nd pair of infralabials contacting each other behind mental scale in D. variegata, although this number is variable in D. indica and D. albifrons); 1st to 4th infralabials in contact with 1st chinshield on each side (vs. usually 2nd to 5th, but less frequently also 6th infralabials in contact with 1st chinshield in D. indica, D. variegata, and D. albifrons); 100–110 subcaudals (vs. 76–93 in D. variegata and 70–96 in D. albifrons; Passos et al., 2005), although this character overlaps with D. indica (Peters, 1960; Hoge and Romano, 1975; Porto, 1993); 15 dorsal scale rows at mid-body (vs. 13 in D. indica), while D. variegata and D. albifrons also exhibit 15 rows; dorsal pattern of strongly black-bordered, straight-sided, upward pointed, light centered triangles with blank and light brown interspaces. The dorsal pattern of D. indica comprises subtriangular, trapezoidal to rounded dark gray-brown to light brown filled blotches or saddles, not bordered by thick straight-sided borders, with interspaces being blank or exhibiting a dark spot (small or none in some specimens in southern Bahia) and paraventral white or yellow blotch, while the dorsal patterns of D. variegata from southern Bahia and D. albifrons are composed of dorsal blotches/saddles vertically stretched rhomboids (more common in D. variegata than in D. albifrons) or bars (less common in D. variegata than in D. albifrons) often with a light center (more common in D. variegata than in D. albifrons) and border zig-zag shaped following the outline of adjacent dorsals, with variable spots in interspaces in D. variegata, rarely in D. albifrons; morphs of Dipsas spp. with straight-sided triangular blotches are rare, but occur in one D. variegata, MNRJ 22771, and more often in allopatric populations of D. variegata from Venezuela (Lotzkat et al., 2008, Natera-Mumaw et al., 2015); cephalic color pattern of D. bothropoides sp. nov. yellow to brown with 3–7 large, dark, light-bordered blotches arranged symmetrically on the pileus (vs. in D. indica: gray to brown with greater number of dark variably sized, yellow-bordered pileus spots not symmetrically arranged except for two parietal blotches; in D. albifrons: unspotted light-colored head with two parallel dark brown blotches extending from parietals onto the neck, occasionally inter-preorbital blotch and brown frontal/parietals; and in D. variegata: variable yellow or brown background, no pileus blotches); eye black (vs. lighter in D. variegata or D. albifrons, and in local D. indica, though the latter species contains some darker morphs with dark eyes, but not as black as in D. bothropoides); ventral pattern consisting of many small square spots (vs. small dots, speckling, or mottled white becoming darker posteriorly in some D. indica, D. albifrons, and D. variegata). Moreover, regional (southern Bahia) D. indica usually possesses a yellow margin (reduced to a few dots or none in some specimens), predominantly along scales contiguous with the eye but also on other cephalic scales, unlike other regional Dipsas species. Dipsas bothropoides sp. nov. has a nuchal blotch < 4 vertebral scales, as in D. variegata or D. albifrons (but > 4 vertebral scales in D. indica), that reaches the rictus, as in most regional D. indica from Bahia, also absent in D. variegata or D. albifrons. 
CommentHabitat: Montane Atlantic rainforest. Note that the paratype is from near sea level and the is holotype from more inland mountains at 700–1,000 m elevation.

Mimicry: T. bothropoides may be a mimic of Bothrops pirajai or Bothrops jararaca given the similar color pattern in these species. 
EtymologyThe specific epithet represents the Latinized form of “bothros” derived from the Greek (ßóθρς), referring to the facial pit, and also referring to the genus Bothrops, the species-rich terrestrial Neotropical pitvipers. The suffix -oides means ‘similar to’ or ‘having the nature of,’ in reference to the great similarity of the dorsal color pattern with many members of the genus Bothrops, especially the sympatric B. jararaca and B. pirajai. 
References
  • Mebert K., Passos P., Fernandes D.S., Entiauspe-Neto O.M., Alvez F.Q., Machado A.S., Lopes R.T. 2020. A new species of snail-eating snake, Dipsas Cope, 1860 (Serpentes: Colubridae: Dipsadinae), from the Atlantic Forest of Brazil. South American Journal of Herpetology 17 (1): 43–62 - get paper here
 
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