Hebius sauteri (Boulenger, 1909)

Li, Mao-Liang, Ren, Jin-Long, Huang, Jun-Jie, Lyu, Zhi-Tong, Qi, Shuo, Jiang, Ke, Wang, Ying-Yong & Li, Jia-Tang, 2022, On the validity of Hebius sauteri maximus (Malnate, 1962) (Squamata, Natricidae), with the redescription of H. maximus comb. nov. and H. sauteri (Boulenger, 1909), Herpetozoa 35, pp. 265-282 : 265

Hebius sauteri (Boulenger, 1909)

Figs 2 , 3 , 4 , 5 , 6

Tropidonotus sauteri Boulenger, 1909, Ann. Mag. Nat. Hist., London. ser.8, 4, 495. Type locality: Kosempo (= Chiahsien, Kaohsiung County), Taiwan, China.

Natrix sauteri - Mell 1931 [1929]: 204; Moichiro 1931: 30; Bourret 1934a: 4; Bourret 1935: 2; Pope 1935: 125 (in part); Smith 1943: 287.

Amphiesma sauteri - Malnate 1960: 51; Zhao and Adler 1993: 228 (in part); Xie et al. 1998: 82; Zhao et al. 1998: 80 (in part); Orlov et al. 2000: 72; David et al. 2007: 60 (in part); Nguyen et al. 2009: 357; Xiang et al. 2012: 230; Wallach et al. 2014: 33 (in part).

Amphiesma sauteri sauteri - Malnate 1962: 272.

Amphiesma sauteri bourreti - Malnate 1962: 273.

Natrix copei Van Denburgh 1909, Proc. California Acad. Sci. 3(3): 52-53. Type locality: "Kanshirei, Formosa" (= Guanziling, Tainan County, Taiwan, China)

Types.

Syntypes. NHMUK (formerly BMNH) 1946.1.13.38-39, a male and a female.

Referred specimens.

19 specimens of H. s. sauteri were examined in this study: CHINA (n = 19) - Taiwan Prov. CAS 18984, Pingtung County; CAS 18988 (paratype of Natrix copei Van Denburgh, 1909), Tainan City, Guanziling. - Fujian Prov. CIB 8476-8479, CIB 78046, Dehua County, Daiyun; CIB 118517, Dehua County, Xinyong Village; CIB 8480, Wuyishan City, Xingcun Township. - Anhui Prov. CIB 8483, Huangshan City. - Jiangxi Prov. SYS r000323, Jinggangshan City, Mt. Jinggang; SYS r001258, SYS r001266, Longnan County, Jiulianshan. - Hainan Prov. YBU 17001, Ledong Li Autonomous County, Jianfengling; CIB 8481. - Guangdong Prov. SYS r001150, Xinyi City, Sihe Township; CIB 118516, Dongguan City, Yinpingshan. - Guizhou Prov. SYS r000275, Libo County, Maolan National Natural Reserve. - Hunan Prov. SYS r001766, Guidong County, Bamianshan.

Diagnosis.

Hebius sauteri can be distinguished from its congeners by the following features: (1) body size small to medium, maximum ToL 455 mm; (2) dorsal scales rows 17-17-17, all weekly keeled or smooth on outmost one or two rows; (3) ventral scales 116-133, subcaudals 60-86, paired; (4) maxillary teeth 23-27, last two or three distinctly enlarged teeth, without diastema; (5) preocular single, occasionally two; (6) supralabials 6-8, 2nd or 2nd-3rd border the loreal; (7) infralabials 6-9, mostly edged with black on their posterior border; (8) single white to orange stripe extends from angulus oris to neck; (9) dorsal body brown to reddish brown, with or without single series of white spots or short band on lateral.

Description.

Head moderately distinct from neck; body size small to medium, ToL 145-455 mm (282-370 mm in males, 145-455 mm in females), TL/ToL 0.24-0.33 (0.27-0.31 in males, 0.24-0.33 in females). Eye large, pupil round.

Dentition. Maxillary teeth 23-27 without diastema, gradually enlarged posteriorly, last two or three distinctly enlarged.

Body scalation. Body scale in 17-17-17 rows, weekly keeled except outer 1-4 rows smooth anteriorly, outer 0-2 rows smooth at midbody and outer 0-1 row smooth posteriorly; ventrals 116-133 (125-130 in males, 116-133 in females); subcaudals 60-86 (60-86 in females, 65-84 in males), paired.

Head scalation. Rostral broad, visible from above; internasals paired, nearly triangular, curved outwards, not in contact with loreal; prefrontals paired, nearly in a diamond shape, wider than long, border on loreal laterally, preocular and supraocular posteriorly; frontal hexagonal, longer than wide, embedded into parietal posteriorly; parietal paired, much longer than wide; nasals wider than high, divided below nostril, borders 1st-2nd supralabials; loreal single, rarely divided (right side of CIB 8481), borders 2nd or 2nd-3rd supralabials, not entering orbit; preocular 1-2; supraocular single, narrowed anteriorly, much wider than high, not in contact with parietal; postoculars 2-3, rarely 4 (left side of CIB 8483); supralabials 6-8, 5th, 6th or 7th highest, 3rd -4th entering orbit, rarely 2nd-3rd (right side of CIB 8480), 3rd only (both sides of CIB 8479) or 4th-5th (left side of CIB 8477); infralabials 6-9, first pair in contact behind the mental, IfL 1-3, 1-4 or 1-5 border the anterior chin shields; temporals variable, in two or three rows, 1-2 + 1-3 + 0/2-3; chin shields paired, posterior pair longer (Figs 3 , 5 , Table 4 ).

Hemipenis. The description of the hemipenis is based on an adult specimen (SYS r000323) from Mt. Jinggang, Jiangxi, southeastern China. Left side of hemipenis fully everted, right side remained in situ for description of retracted condition.

The everted hemipenis is thin and short, Y-shaped, shallowly bilobed. Both sulcate and asulcate surfaces densely ornamented with spinules, a large basal hook present at proximal part. Sulcus spermaticus single, extending to base of inner right lobe where it takes a centripetal position. Sulcus lip highly developed and raised, walls covered with spinules (Fig. 6 ).

In situ hemipenis extends to 7thSC with its crotch extending to 6thSC; crotch of m. retractor penis magnus extending to 8thSC, origin of m. retractor penis magnus at level of 20thSC.

Coloration in life.

Based on two adult female specimens from Guangdong (CIB 118516) and Fujian (CIB 118517), China. Dorsal surface of the head reddish-brown with irregular grayish-green spots, a pair of elliptic orangish spots symmetrically present on inner sides of parietals, just separated by parietal sutures; ventral side of head whitish; 1st-5th supralabials white with black edges posteriorly, 6th or 6-7th brown on their upper part; infralabials white with black edge between each scale; an orange stripe extends from angulus oris to dorsal side of neck, 1-2 scales wide; dorsal surface of body and tail dark brown to reddish-brown, top with or without an ill-defined dark olive stripe scattered with black spots extends from neck to the end, a reddish-brown lateral streak on 4th-6th body scale rows extends from neck to tail with series of orange spots separated by 2-3 scales above; ventral and subcaudal sides cream white, with black spots on both ends of the ventral scale forming a chain-like pattern on the ventral view; sclera brown (Fig. 4 ).

Coloration in preservative.

Largely the same as in life except dorsal side of head brown to dark brown, scattered with small black spots; ventral side of head cream white; a pale yellow stripe extends from angulus oris to dorsal side of neck, 1-2 scales wide; dorsal of body and tail brown to dark brown, a lighter lateral streak on 4th-6th body scale rows presents or not, covered with series of cream white spots separated by 2-3 scales; ventral and subcaudal sides cream to pale white; sclera black (Figs 3 , 5 ).

Etymology.

Hebius sauteri was named after the collector of the type specimens, Hans Sauter (1871-1943), a German naturalist ( Boulenger 1909). We suggest that the Chinese common name of this species remains unchanged, as "Zōng Hēi Fù Liàn Shé ( 棕黑腹链蛇)”.

Natural history.

This species inhabits flatlands and mountain regions up to 1,500 m and prefers terrestrial microhabitat such as grassland, woodland, and bushes near water bodies; diurnal, occasionally found at night, preys include earthworms, slugs, snails and tadpoles; oviparous (this study; Pope 1935; Zhao 2006; Xiang et al. 2009).

Distribution.

Hebius sauteri is currently known to be reported in Vietnam (Ngan Son, Lang Son, Tam Dao and Ba Vi), and China, including Taiwan (Kaohsiung County [type locality]), Fujian (Dehua County), Anhui (Huangshan County), Jiangxi (Jinggangshan City, Longnan County), Hunan (Yizhang County), Guangdong (Lianping County, Xinyi County, Deqing County, Ruyuan County, Dongguan City), Guangxi (Jinxiu County, Quanzhou County, Rongxian County, Beiliu County, Long’an County), Guizhou (Libo County, Leishan County), Yunnan (Jinping County), Hainan (Qiongzhong County, Wuzhishan County, Baisha County, Lingshui County) ( Deuve 1970; Wu 1985; Zhao 2006; Yang et al. 2008; Nguyen et al. 2009; Zhang 2009; Li 2011; Shen 2014; this study) (Fig. 2 ).

Remarks.

Natrix copei (Van Denburgh, 1909) was described by Van Denburgh (1909) based on specimen form “Kanshirei” (now Guanziling), Tainan County, Taiwan Province, China, the description of this species was published just 20 days after that of Tropidonotus sauteri by Boulenger (1909) (now Hebius sauteri ) ( Stejneger 1910), Stejneger (1910) considered Natrix copei a synonym of Natrix sauteri . We examined the paratype of Natrix copei , and its morphological characters are in the range of variation of H. sauteri , which also supports the treatment of Natrix copei as an objective synonym by Stejneger (1910). The taxonomic status of the subspecies H. s. bourreti is discussed below.

Figures

Figure 2. Distribution of Hebius maximus comb. nov. and re-defined distribution of H. sauteri in China based on museum collections, previous studies, and this study (Triangle: Hebius maximus comb. nov., round: H. sauteri sauteri, square: H. sauteri bourreti. Type localities are colored red).

Figure 3. Dorsal (left), lateral (middle) and ventral (right) of the head comparisons between Hebius sauteri and Hebius maximus comb. nov. Row D photographed by Kai Wang, others were photographed by Jun-Jie Huang. A-C. Hebius maximus comb. nov. (A. CIB 118635, Mt. Laojun, Pingshan County, Sichuan, China; B. CIB 8482, Bijie City, Guizhou, China; C. CIB 8458, Yingjing County, Sichuan, China) D-F. H. sauteri (CAS 18984, Pingtung County, Taiwan, China; CIB 118516, Yinpingshan, Dongguan City, Guangdong, China; CIB 118517, Daiyun, Dehua County, Fujian, China).

Figure 4. Hebius sauteri. A. CIB 118516, Yinpingshan, Dongguan City, Guangdong, China; B. CIB 118517, Xinyong Village, Dehua County, Fujian, China. Photographs by Jin-Long Ren.

Figure 5. Preserved specimen of H. sauteri (CAS 18988), from the type locality of the species, Guanziling, Tainan City, Taiwan, China. Not to scale. Photographs by Kai Wang.

Figure 6. Photographs of the hemipenis of H. sauteri (SYS r 000323, Mt. Jinggang, Jiangxi Province, China), photos from left to right showing sulcate, asulcate, and apical sides of the hemipenis. Photographs by Jun-Jie Huang.