Bothrops sonene, Carrasco & Grazziotin & Farfán & Koch & Ochoa & Scrocchi & Leynaud & Chaparro, 2019

Carrasco, Paola A., Grazziotin, Felipe G., Farfán, Roy Santa Cruz, Koch, Claudia, Ochoa, José Antonio, Scrocchi, Gustavo J., Leynaud, Gerardo C. & Chaparro, Juan C., 2019, A new species of Bothrops (Serpentes: Viperidae: Crotalinae) from Pampas del Heath, southeastern Peru, with comments on the systematics of the Bothrops neuwiedi species group, Zootaxa 4565 (3), pp. 301-344 : 310-318

Bothrops sonene sp. nov.

Figures 4 –7, Table 1

Bothrops gr. neuwiedi: Cadle et al. 2002: 102 (MUSM 33453, from Pampa Juliaca, department of Madre de Dios, Peru).

Holotype. An adult female, MUBI 12319 , collected on April 23, 2013 by J.A. Ochoa at Pampa Juliaca (12°57’46.79” S, 68°55’22.61” W; 206 m above sea level, asl hereafter), Pampas del Heath ( Bahuaja-Sonene National Park ), district of Tambopata, province of Tambopata, department of Madre de Dios, Peru ( Figs. 5A, 5E , 6B ).

Paratypes. Three specimens, all from Pampa Juliaca , Pampas del Heath ( Bahuaja-Sonene National Park ), district of Tambopata , province of Tambopata, department of Madre de Dios, Peru. An adult male, MUBI 14679 , collected on August 14, 2015 by J.A. Ochoa, G. Sebastián & K. Huamani (12°57’47.47” S, 68°54’57.01” W; 210 m asl) ( Figs. 5B, 5F , 6C ). An immature male, MUSA 4350 , collected on November 13, 2015 by R. Santa Cruz (12°57’2.47” S, 68°54’51.05” W; 209 m asl) ( Figs. 4 , 5C , 6A, D ). An immature male, MUSM 33453 , collected on between June 02–25, 1996 by J. Cadle (12°59’20.10” S, 68°55’48.45” W; 208 m asl) ( Fig. 5D ) .

Diagnosis. Bothrops sonene can be distinguished from its congeners by the exclusive combination of the following morphological characters: dorsal color pattern of body consisting of small C-shaped blotches, apparently unique for the genus; postocular stripe originating posteriorly to the eye, two or two and a half scales width, covering three or four posterior supralabials; dorsum of the head with paired markings arranged symmetrically; venter cream heavily speckled with brown; lacunolabial scale absent, prelacunal discrete contacting second supralabial; prefoveals three to five; subfoveal single and usually present; postfoveals absent to two; canthals two; intersupraoculars seven; suboculars one or two; postoculars two or three; supralabials seven or eight; infralabials nine to eleven; interrictals 26–27; middorsal scales 23–25; ventrals 172 in the female, 169–173 in males; subcaudals divided, 45 in the female, 50 in males.

Comparisons. Bothrops sonene is easily distinguished from its congeners by its unique dorsal color pattern of relatively small and C-shaped blotches. It is distinguished from B. atrox , B. barnetti , B. jararaca , B. jararacussu , B. pictus and B. taeniatus by the absence of a lacunolabial scale (= prelacunal fused with second supralabial), a condition shared with the species of the B. alternatus and B. neuwiedi groups. Similar C-shaped (or in the shape of a headphone) dorsal blotches are present in B. alternatus , B. cotiara , B. fonsecai , B. itapetiningae and B. jonathani ( B. alternatus group excluding B. ammodytoides , whose dorsal blotches are quadrangular); however, the blotches of B. sonene are smaller and rounder than those species, the lower set of blotches are well separated from each other (except in B. itapetiningae ), and blotches are bordered with black (vs. bordered with white). The new species can also be distinguished from some species of the B. alternatus group by the postocular stripe starting posteriorly to the eye (vs. below the eye in B. alternatus , B. jonathani and B. ammodytoides ), number of prefoveals (3–5 vs. 3– 15), subfoveals (0–1 vs. 1–6), supralabials (8 vs. 8–12) and infralabials (9–11 vs. 10–15), length of hemipenes (relatively short vs. long), and fusiform and divergent hemipenial lobes (vs. parallel and subcylindrical lobes). Bothrops sonene is more similar to the B. neuwiedi group of species than to the rest of the species of the genus. They all have a symmetrical pattern of blotches in the dorsum of the head, a brown and relatively wide postocular stripe that extends posterior to the rictal region, a discrete prelacunal scale contacting the second supralabial, and similar number of prefoveals, subfoveals, supralabials and infralabials. They also bear resemblance in hemipenial morphology; the overall morphology of the hemipenis of the new species is identical to that of the species of the B. neuwiedi group, which is characteristic of that group. Differences in coloration and scalation between Bothrops sonene and the species of the B. neuwiedi group are summarized in Table 2.

Description of the holotype. Adult female; snout-vent length 950 mm; tail length 123 mm; caudal spine length 8.9 mm; head length 44.8 mm; maximum head width 36.5 mm; interorbital space 7.2 mm; internasal space 2.4 mm; dorsal scale rows 23–24–19; ventral scales 172; subcaudal scales 45/45; rostral trapezoidal; canthus rostralis elevated forming a ridge; internasals 1/1; canthals 1/1, similar size than internasals; intercanthals, keeled, 4–5–5; intersupraoculars, keeled, 7; scales around supraoculars 11/10; supralabials 8/7; infralabials 10/11; postnasal in contact (left side) or not in contact (right side) with first supralabial; loreal subtriangular; prefoveals 5/ 5; subfoveal 1/0; postfoveals 1/1; prelacunal contacting the second supralabial; sublacunals 1/1; supralacunals 1/1; suboculars 2/1; postoculars 3/2; upper preoculars contributing to the canthus rostralis, right preocular divided; upper and medial preoculars in contact with the orbit; inferior preoculars not in contact with the orbit; scales between suboculars and 3 rd –4 th supralabials 2/2; scales between suboculars and 4 th supralabial 2/2; scales between suboculars and 4 th –5 th supralabials 2/2; interrictals 26; gulars 4. Dorsal background color of body light brown; dorsum with dark brown, black-edged blotches, each blotch composed of dorsal and ventral C-shaped forms; ventral surface of body cream, heavily speckled with dark brown spots; dorsum of head with symmetrically arranged brown and dark-edged blotches, including a single blotch on the snout, a pair of blotches near supraoculars, a pair of parietal stripes that extend posterior-laterally, and a pair of occipital stripes that extend posteriorly; head with lateral, brown and black-bordered postocular stripes originating posterior to each eye.

Color pattern variation. Dorsal background color of head and body light brown with olive green tonality in live specimens; dorsum of body with 28–31 C-shaped, dark brown and black-edged, dorsal blotches; ventral portion of blotches formed by a pair of mostly-fused round blotches, forming a single curved blotch; ventral surface of body cream, speckled with dark brown spots, the spots forming a transversal stripe in some ventral scales; dorsum of head with brown, dark-edged, symmetrically arranged blotches; postocular stripes extend 1.5–3 scales posterior to rictal region, encroaching 3–4 supralabials and 1–2 infralabials; supralabials with dark brown spots, more conspicuous between 3 rd –5 th supralabials; infralabials mostly bordered with dark brown pigments. Gular region of males cream with irregular dark brown markings, absent in the female of the type-series suggesting a putative sexual dimorphism in this character; tip of tail pale ventrally in adult female and male (MUBI 14679); both dorsal and ventral surface of tip of tail pale in subadult males (MUSA 4350, MUSM 33453).

Hemipenis morphology. Based on MUBI 14679 and MUSA 4350. Organ strongly bilobed; hemipenial lobes fusiform and divergent, 67% of total hemipenial length; hemipenial body 33% of total hemipenial length; capitulum longer on sulcate side, occupying 62% of lobe; base of capitulum covered with spinulate calyces; hookshaped spines distributed asymmetrically on lobes, big sized spines located proximally; small, thin and curved spines present in the intralobular region, located distally on the lobes; hemipenial body covered with spinules; microornamentation on intrasulcar region absent; sulcus spermaticus bifurcating proximal to capitulum and ornamented with spinules (Fig. 7).

Distribution and natural history. Known to occur in Pampas del Heath, in the Bahuaja-Sonene National Park, province of Tambopata, department of Madre de Dios, Peru, at between 206–210 m asl ( Fig. 2 ). The area is situated in southeastern Peru, near the border with Bolivia, between the Tambopata and Heath Rivers. Bothrops sonene is known from four close localities situated inside of the savannas of the Peruvian Pampas del Heath.

The specimens were collected at the end of the rainy season and during the dry season. They have diurnal activity; during the day they were observed on the ground and above grass at 500 mm above ground, sometimes slithering rapidly. They are associated with water bodies, which are widespread along the savannas of Pampas del Heath in rainy seasons (the soil is flooded up to 250 mm from the ground) and restricted to the “Aguajales” (Palm Tree areas) in dry seasons (Fig. 8).

Etymology. The specific epithet “sonene” is derived from the tacana language, and is the name used by the Ese’ejja for the Heath River. The Ese’ejja are an Amazonian community that inhabits the areas surrounding the Madre de Dios, Tambopata, and Heath Rivers, in the border between Peru and Bolivia. We named the new species “sonene” because its type locality, the Peruvian Pampas del Heath, is located near the Heath River.

B. sonene B. cf. sonene B. neuwiedi B. aff. neuwiedi B. mattogrossensis B. aff. B. aff. B. aff. mattogrossensis 1 mattogrossensis 2 mattogrossensis 3

BP C-shaped blotches Trapezoidal or C- Trapezoidal or Trapezoidal or Trapezoidal Trapezoidal Trapezoidal Trapezoidal shaped blotches subtriangular subtriangular blotches blotches blotches blotches blotches blotches

PM Stripes Stripes or rounded Stripes Stripes Rounded blotches Rounded blotches Rounded blotches Rounded blotches blotches

POSw 2–2.5 2 – 3 2.5 – 3 1.5 – 3 2 – 2.5 2 – 3 1.75 – 3 2 – 2.5 x=2.2 x=2.4 x=2.8 x=2.3 x=2.2 x=2.2 x=2.2

SLPOS 3–4 3 – 4 2 2 – 3 3 – 4 2 – 4 2 – 4 2 – 3 x =3.2 x=3.3 x= 2 x =3.8 x= 3 x = 3 x =2.7

V 164–172 173 – 178 166 – 172 172 – 184 183 – 190 163 – 173 155 – 165 167 – 177 x=169.5 x=175 x=169 x=177.9 x=186.5 x=167.5 x=160 x=173

MD 23–25 25 – 27 25 – 27 24 – 28 23 – 25 21 – 25 21 – 23 21 – 25 x = 24 x = 26 x =26.3 x=25.7 x=23.5 x= 23 x =22.6 x=22.8

SL 8 8 – 9 8 – 11 8 – 9 7 – 10 7 – 8 7 – 8 7 – 9 x = 8 x = 9 x =8.4 x=8.3 x=7.6 x=7.6 x=8

IL 10–11 10 – 11 11 – 13 10 – 13 11 – 12 9 – 11 8 – 10 9 – 12 x=10.5 x= 10 x =11.7 x= 11 x =11.5 x= 10 x =9.4 x=10.8

ICm 4–6 4 – 6 6 4 – 6 3 – 4 3 – 5 3 – 5 3 – 7 x =5.2 x=5.2 x= 5 x =3.7 x=3.7 x=4.2 x=5

ISPO 7 7 8 – 11 7 – 11 5 – 8 5 – 7 4 – 7 6 – 8 x =9.3 x= 8 x = 7 x =6.3 x=5.6 x=7.3

PF 3–5 4 5 – 7 2 – 6 3 – 6 2 – 5 2 – 4 4 – 5 x = 4 x =6.3 x=3.9 x=4.5 x=3.2 x=3.2 x=4.6

SBF 1 0 – 1 1 – 3 0 – 2 0 – 2 0 – 1 0 – 1 1 – 3 x =0.3 x= 2 x = 1 x =0.8 x=0.5 x=0.6 x=1.6

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TABLE 2. (Continued)

TABLE 2 . Continues .

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TABLE 2. (Continued)

Figures

FIGURE 2. Distribution of the lineages of the Bothrops neuwiedi species group. Symbols are as follow: stars = type localities, circles = records of specimens with DNA samples, squares = records of specimens that were only examined morphologically.

FIGURE 4. Bothrops sonene sp. nov. (MUSA 4350) from Pampas del Heath, Bahuaja-Sonene National Park, department of Madre de Dios, Peru.

FIGURE 5. Dorsal view of the holotype (MUBI 12319—A, 1073 mm TL) and paratypes (MUBI 14679—B, 553 mm TL, MUSA 4350—C, 435 mm TL, MUSM 33453—D, 339 mm TL) of Bothrops sonene. Ventral view of holotype (MUBI 12319— E) and paratype (MUBI 14679—F). TL= total length.

FIGURE 6. Dorsolateral view of head of the paratype (MUSA 4350—A, D, 20 mm HL) and holotype (MUBI 12319—B, 44.8 mm HL), and dorsal view of head (MUBI 14679—C, 23 mm HL) of the paratype of Bothrops sonene. HL= head length.