A new species of pitviper of the genus Protobothrops from China (Squamata: Viperidae) Author Yang, Jian-Huan Author Orlov, Nikolai L. Author Wang, Ying-Yong text Zootaxa 2011 2936 59 68 journal article 46497 10.5281/zenodo.278107 572f25c8-cd28-425b-a7cc-2df8744fc69e 1175-5326 278107 Protobothrops maolanensis sp. nov. ( Figs. 1–3 , Table 1 ) Holotype . SYS r000211, adult male ( Fig. 1 ) from Maolan National Nature Reserve, Maolan town, Libo County, Guizhou, China ( 25°12'50.0'' N , 107°59'56.04'' E , 560 m a.s.l.), collected by Sheng Zheng and Jian-Huan Yang on 18 May 2010 . Paratypes . SYS r000210, adult male, same data as holotype . SYS r000276, adult male and SYS r000277, adult female, from the same locality as holotype , collected by Jian-Huan Yang, Yun Li and Run-Lin Li on 10 August 2010 . Diagnosis. Protobothrops maolanensis sp. nov. differs from other species of Protobothrops by the combination of the following characters: (1) a relatively small body size (TL 609–805 mm ); (2) dorsal scales keeled throughout, in 23(rarely 25): 21(19): 17(15) rows; (3) ventral scales 186–193 in male, 190 in female; (4) subcaudals 78–85 pairs in male, 74 in female; (5) 7–8 supralabials; (6) 11–12 infralabials; (7) 10–12 scales between midsection of supraoculars in a line; (8) the 4th supralabials separated from suboculars by one row of scales; (9) 68–72 small and thin body bands across body and tail; (10) TaL/SVL ratio 0.238–0.269 in male, 0.190 in female; (11) SL/ HL ratio 0.308–0.326 in male, 0.344 in female; (12) ED/HL ratio 0.164–0.176 in male, 0.152 in female; and (13) the unique color pattern. See below for differential diagnosis by way of comparison with all other species in the genus. FIGURE 1. A–E (Holotype of Protobothrops maolanensis sp. nov. , SYSr000211). A: General aspect in life; B: dorsal views of head; C: ventral views of head; D: lateral views of head; E: dorsal view of midbody (showing thin dark brown bands and keeled scales). F (Paratype, SYS r000210): left hemipenis in asulcate view. Photographs by J.H. Yang. FIGURE 2. A: paratype of Protobothrops maolanensis sp. nov . (female, SYS r000277); B: holotype of P. trungkhanhensis (male); C: paratype of P. trungkhanhensis (female); D: karst forest in Maolan National Nature Reserve; E: microhabitats of the type locality. Photographs by Sheng Zheng (A), N. L. Orlov (B, C) and J. H. Yang (D, E). Description of holotype . Adult male with detailed measurements as listed in Table 1 . Body moderately elongate, thin and slightly compressed; head elongate and triangular, distinctly wider than neck, covered with very small, convex and irregular shaped scales, 0.69 times wider than long; upper head scales smooth anteriorly, keeled on occipital, 10 scales between midsection of supraoculars in a transverse line; snout elongate, SL/HL ratio 0.31, nearly twice ED; eye convex, pupil vertical. TABLE 1. Measurements and scale characters of the type specimens of Protobothrops maolanensis sp. nov. See text for abbreviations.

SYS r000211 Holotype	SYS r000210 Paratype	SYS r000276 Paratype	SYS r000277 Paratype
Sex SVL TaL TaL/SVL	male 634 171 0.27	male 638 160 0.25	male 492 117 0.24	female 521 99 0.19
TL HW	805 16.6	798 16.6	609 13.8	620 14.8
HL HW/HL SL SL/HL	24.0 0.69 7.7 0.32	24.5 0.68 7.5 0.31	19.6 0.70 6.4 0.33	21.7 0.68 7.5 0.35
ED ED/HL	4.2 0.18	4.3 0.18	3.2 0.16	3.3 0.15
EN supralabials	4.8 8/8	4.9 7/8	4.2 8/8	4.8 8/8

3rd supralabials contact subocular infralabials	no 11/12	yes 11/12	yes 12/12	no 11/11
SBSO	10	11	12	10
L canthal scales	2 2/2	2 3/3	2 3/3	2 4/4
pre-V V Scd Sq1: Sq2: Sq3	2 186 85 25: 21: 17	2 188 78 23: 19: 15	1 193 80 23: 21: 17	2 190 74 23: 21: 17
outermost row of dorsal scales	strongly keeled	strongly keeled	strongly keeled	smooth anteriorly, weakly keeled posteriorly
body bands	71	72	68	72

Rostral triangular, about 1.3 times broader than high, barely visible from above, broader posteriorly with three apical scales bordered laterally by internasals; internasals not in contact with rostral and separated from each other by four small scales; nasal trapezoid, undivided, with a round nostril opening in middle; 2/2 (L and R, hereafter) canthal scales between supraocular and internasal, distinctly larger than adjacent snout scales, anterior one largest, bordering sharp, raised canthus rostralis ; two loreals, approximately equal size; supraocular large and elongate, wider than adjacent head scales, pointed anteriorly and 0.58 times as wide as interspace between supraoculars and surrounded by 13/13 small scales; two elongate upper-preoculars above loreal pit in contact with posterior loreal, lower one longer than upper and forming upper margin of loreal pit; one elongate lower-preocular, forming lower margin of loreal pit; two small postoculars; one elongate, thin, crescent-shaped subocular, separated from lower preocular by one scale; temporals numerous, upper ones keeled, lower ones smooth, those bordering supralabials larger than latter; anterior part of nasal large, turning over canthus rostralis and visible above; 8/8 supralabials, 2nd forming anterior margin of loreal pit and separated from nasal by one small scale, 3rd largest, 3rd and 4th separated from subocular by a row of scales on both sides; 11/12 infralabials, 1st pair in contact with each other, first three in contact with anterior chin shield; posterior chin shield small, scarcely differentiated from postgenials. Dorsal scales narrow, pointed and strongly keeled throughout; in 25 rows on anterior body, 21 rows at midbody, 17 rows at posterior of body; pre-ventral scales two; ventral scales 186; anal scale undivided; subcaudals divided, 85 pairs. Coloration in life. Grayish brown above, with 71 dark brown transverse bands with discontinuous pale yellow rough edges, across body and tail. The dark brown bands are relatively small and thin, occupy about six dorsal scale rows in width and 0.5–1.5 dorsal scales in length and often broken into spots. The bands become gradually closer towards tail; tip of tail dark brown with scatted pale spots. Underside gray, with a series of pale brown blotches on both sides. Dorsal head gray brown, with indistinctly symmetrical dark streaks. A postorbital thin dark brown stripe with discontinuous pale yellow edge, starting from posterior margin of eye, extending across upper temporals posteriorly, ending above corner of lip. Variation. Measurements and scale characters of all the specimens are listed in Table 1 . All the paratypes match the holotype in overall morphology and scalation. Dorsal scales in 23 rows on anterior part of body in all three paratypes , 19 rows on mid-body in SYS r000210, 15 rows on posterior part of body in SYS r000210; 3rd supralabials touching the subocular in SYSr000210; 3/3 canthal scales in SYS r000210 and 0 0 0 276, 4/ 4 in SYS r000277. Sexual dimorphism. No significant sexual dimorphism occurs, other than the female having fewer subcaudals than the males (74 vs. 78–85) and the outermost row of dorsal scales are weakly keeled in the female, nearly invisible at anterior body, while the dorsal scales are strongly keeled throughout in male. Hemipenis. The everted left hemipenis ( paratype , SYS r000210) is 18.5 mm long, robust and forked, spinous proximally and calyculate distally; it bears two lobes covered with spines and microspines, extending to the 8th subcaudal. Sulcus spermaticus forked at the truncus 6.5 mm from the base of the hemipenis. Strong spines on asulcate side at the base part of each lobe, followed by smaller spines more distally; sulcate side covered with microspines. The hemipenis of P. maolanensis closely resembles the Type 3 spinose hemipenis of Malhotra & Thorpe (2004). FIGURE 3. The type locality of Protobothrops maolanensis sp. nov. , Maolan National Nature Reserve (circle, “MNNR”), Maolan town, Guizhou, China. Distribution and natural history. Currently, P. maolanensis is known only from the type locality, Maolan National Natural Reserve, Maolan town, Guizhou, China . All individuals were collected at 550–600 m elevation in a subtropical karst evergreen forest. They were found on the ground under scrub at night; no individuals were observed during the day. Other species of amphibians and reptiles at the same habitat in the type locality during the survey included Polypedates megacephalus (SYS a000959-962), Kurixalus odontotarsus (SYS a000926-931), Microhyla fissipes (SYS a000940), Microhyla heymonsi (SYS a000938-939), Goniurosaurus cf. luii (SYS r000217-219), Takydromus sp., Sphenomorphus cf. indicus , Protobothrops mucrosquamatus (SYS r000238), Dinodon flavozonatum (SYS r000274), Achalinus spinalis (SYS r000273) and Bungarus multicinctus (SYS r000305). Etymology. The specific epithet refers to the type locality, Maolan Town, Guizhou, China . We propose the English name “Mao-lan pitviper”. Comparisons. Protobothrops maolanensis sp. nov. can be readily distinguished from P. cornutus and P. sieversorum by the absence of horn-shaped projections on the supraocular and a different color pattern. Protobothrops maolanensis sp. nov. differs from P. elegans by having fewer mid-body dorsal scales (19–21 vs. 23–25), 4th supralabials separated from suboculars by one row of scales (two in P. elegans ), two loreals (one in P. elegans ), dorsal scales keeled throughout (outermost row of dorsal scales smooth in P. elegans ) and a different color pattern. Protobothrops maolanensis sp. nov. differs from P. flavoviridis by having fewer mid-body dorsal scales (19–21 vs. 33–40), two loreals (one in P. flavoviridis ), lower number of ventral scales (186–193 vs. 222–237 in P. f l a v o - viridis ), dorsal scales keeled throughout (outermost row of dorsal scales smooth in P. flavoviridis ) and a different color pattern. Protobothrops maolanensis sp. nov. differs from the P. jerdonii complex ( P. j. jerdonii , P. j. bourreti , P. j. xanthomelas ) by its smaller size, slender body, dorsal scales keeled throughout (outer one or two rows of dorsal scales smooth in P. jerdonii ) and by a clearly different color pattern. Protobothrops maolanensis sp. nov. differs from P. kaulbacki by its smaller size, fewer mid-body dorsal scales (19–21 versus 25), dorsal scales keeled throughout (outermost row of dorsal scales smooth in P. kaulbacki ), fewer ventral scales (186–193 vs. 201–212 in P. kaulbacki ), and by a clearly different color pattern. Protobothrops maolanensis sp. nov. differs from P. mangshanensis by its smaller size (largest TL 805 mm in P. maolanensis sp. nov. vs. largest TL 2375 mm in P. mangshanensis ), internasals separated by scales (touching each other in mangshanensis ), two loreals (one in mangshanensis ), fewer infralabials, (11–12 vs. 13–16), more subcaudals (74–85 vs. 60–67), fewer mid-body dorsal scales (19–21 vs. 25), dorsal scales keeled throughout (outermost row of dorsal scales smooth in P. mangshanensis ) and a different color pattern. Protobothrops maolanensis sp. nov. differs from P. mucrosquamatus by its smaller size (largest TL 805 mm vs. largest TL 1310 mm in P. mucrosquamatus ), fewer mid-body dorsal scales (19–21 vs. 23–27), fewer ventral scales (186–193 vs. 194–233 in P. mucrosquamatus ), dorsal scales keeled throughout (outermost row of dorsal scales smooth in P. mucrosquamatus ) and a different color pattern. Protobothrops maolanensis sp. nov. differs from P. tokarensis by having fewer mid-body dorsal scales (19–21 vs. 31–32), two loreals (one in P. tokarensis ), fewer ventral scales (186–193 vs. 203–208 in P. tokarensis ), dorsal scales keeled throughout (outermost row of dorsal scales smooth in P. tokarensis ) and a different color pattern. Protobothrops maolanensis sp. nov . differs from P. xiangchengensis by having fewer mid-body dorsal scales (19–21 vs. 25), more subcaudals (74–85 vs. 50–66), dorsal scales keeled throughout (outer two rows of dorsal scales smooth in P. xiangchengensis ), absence of a dark brown blotch under loreal pit (present in P. xiangchengensis ). Protobothrops maolanensis sp. nov. is superficially most similar in appearance to P. trungkhanhensis from Vietnam , from which it differs by having more dorsal scale rows on the anterior part of the body (23–25 vs. 19), outermost row of dorsal scales on body strongly keeled in males and weakly keeled in females (dorsal scales strongly keeled throughout in both male and female P. trungkhanhensis ), lower ED/HL ratio (0.15–0.18 vs. 0.23), higher HW/HL ratio (0.68–0.70 vs. 0.65–0.66), and fewer body bands (68–72 vs. 76–84).