Amphiesma boulengeri (Gressitt, 1937)
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publication ID |
https://doi.org/ 10.11646/zootaxa.3694.4.1 |
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publication LSID |
lsid:zoobank.org:pub:6F2F58C1-3927-4E66-B09C-AFAF4B4A2C71 |
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DOI |
https://doi.org/10.5281/zenodo.6165422 |
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persistent identifier |
https://treatment.plazi.org/id/855087F6-7E1E-FFA7-FF5F-72D9B7FA6303 |
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treatment provided by |
Plazi |
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scientific name |
Amphiesma boulengeri (Gressitt, 1937) |
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Amphiesma boulengeri (Gressitt, 1937)
( Fig. 4 View FIGURE 4 A–B, 5)
“ Natrix khasiensis > gilhodesi [sic]”.—Mell 1931a: 203—Pope 1935: 90 (discussion of the taxon).—Status. None, not a binominal combination.
Natrix boulengeri Gressitt, 1937: 125 . – Type locality. “Tai-yong, alt. 640 meters, E. Kwantung province, south-eastern China (lat. 23° 34' N., long. 115° 55’ E), now Dayang, Jiexi County, Guangdong Province, People’s Republic of China. – Holotype. MVZ 23623 (adult female); collected by J. L. Gressitt, 5 Aug. 1936.
Natrix boulengeri .—Gressitt 1941: 4 & 36, Pl. 1.—Anonymous 1977: 63 & 87—Yang et al. 1983: 43—Hu et al. 1980: 66— Zhong & Wu 1981: 105—Wu et al. 1985: 173, Fig. 62, 398: Pl. 29–2—Wang 1987: 182.
Amphiesma boulengeri .—Malnate 1960: 50, 52 & 57—Tian et al. 1986: 116 & 149—Yang & Inger 1986: 7—Zhao 1986: 239—Kou & Zhang 1987: 364—Welch 1988: 29—Zhao 1991: 358; 1997: 54; 2006: (I)162, (II)82: Fig. 45- 1–45-2 —Su 1992: 199—Zhao & Adler 1993: 225—Zhao & Yang 1997: 201 & 202—Deng & Ye 1998: 92—Huang et al. 1998: 79, Fig. S-23, 4th B&W plate (unnumbered): Fig. 7 View FIGURE 7 —Karsen et al. 1998: 46, 47 & 106—Xie et al. 1998: 82—Ye et al. 1998: 3 & 11—Zhao et al. 1998: 51, 55 & 56: Fig. 18; 2000a: 232; 2000b: 205; 2004: 317—Yang 1999: 65—Zhang 1999a: 430— Zhang 1999b: 65; 2009: 80 & 81, Pl. 4: Fig. 79—Orlov et al. 2000: 71; 2008: 84—Zhong 2000: 6, 58 & 59: Fig. 46a–b; 2004: 223—Iskandar & Colijn 2001: 95—He & Zhou 2002: 167—Ji 2002: 174 & 175—Song et al. 2002: 73—Zhou & Yu 2002: 237—Fellowes et al. 2003a: 11, front cover; 2003b: 23—Nguyen et al. 2005: 74; 2009: 352—David et al. 2007: 41 & 55—Mo et al. 2007: 344—Ziegler et al. 2007: 22 & 37—Grismer et al. 2008: 166 & 173—Yang & Rao 2008: 254 & 372: Fig. 9 (identification uncertain)—Chen et al. 2009: 929—Zeng 2009: 754—Das 2010: 150: Pl. 67: Fig. 4 View FIGURE 4 , 331 (at the exception of the mention of Thailand: Nakhon Si Thammarat Province)—Luo et al. 2010: 73—Teynié & David 2010: 284 & 308—Bain & Hurley 2011: 104 & 129—Li 2011: 158—Shi 2011: 214, Pl. 15: Fig. 81.
Tropidonotus modestus (nec Tropidonotus modestus Günther, 1875 , a valid species)—Werner 1904: 363 & 375 (probably refers to A. boulengeri ; see Pope 1935: 90)—Mell 1922: 116—Vogt 1922: 137—Bourret 1927: 237 (? Probably in part).
Natrix modesta (nec Tropidonotus modestus Günther, 1875 ).—Angel 1929: 76—Mell 1929: 5, 16, 145, 147 & 157; 1931: 203—Bourret 1936a (in part): 103 & 113; 1936b (in part): 72 (text only; Fig. 30 on p. 74 refers to another species; see below); 1940b: 42 (?)—Smith 1943: 290 (in part; specimens cited on p. 291 from “Upper Laos (Chieng Khoung [sic])”, “ Cambodia (Kamchay Mts.)”, “S. Annam (Langbian Plateau)”, and “Pen. Siam ”)—Deuve 1970: 84 & 88 (in part; specimens de “Xieng Khoang”)—Tran et al. 1981: 380.
Amphiesma modesta (nec Tropidonotus modestus Günther, 1875 ).—Campden-Main 1970: 15.—Saint Girons 1972: 80 (in part)—Dao 1981: 7 (?)—Nguyen 1981: 11—Ho & Nguyen 1982: 139—Nguyen et al. 1995a: 32—Daltry & Chheang 2000: 232.—Le et al. 2001: 27—Daltry & Traeholt 2003: Appendix III: page ff.
Amphiesma modestum .—Das 2010: 333 (in part; only for mentions of northern Laos, south-central Cambodia and central Vietnam).
Natrix modesta modesta (nec Tropidonotus modestus Günther, 1875 , a valid species).—Bourret 1936b (in part): 73 (text only; Fig. 30 on p. 74 refers to another species; see below); 1940c (in part): 53.
Tropidonotus johannis (nec Tropidonotus johannis Boulenger, 1908 , a valid species).—Smith 1921: 426 (see Pope 1935: 90)— Bourret 1936a: 27.
Natrix modesta johannis (nec Tropidonotus johannis Boulenger, 1908 ).—Bourret 1936a: 103 (in part) & 113; 1936b: 74 (in part; record from “Sud-Annam (Plateau du Lang Bian)”).
Natrix khasiensis (nec Tropidonotus khasiensis Boulenger, 1890 ).—Mell 1929: 148 (in part)—Bourret 1934a: 149 & 150 [Reprint: 3 & 4]; 1935a: 239 & 240 [Reprint: 5 & 6]; 1935b: 259 & 261 [Reprint: 1 & 2]; 1935c: 289 & 290 [Reprint: 1 & 2]; 1936a: 32, 89: Fig. a, 103 & 113; 1936b: 69, Fig. 28 (in part for the text; figure depicting Amphiesma boulengeri ); 1937a: 28; 1937b: 58 & 71; 1940a: 23; 1940b: 42, 44 & 47; 1940c: 53; 1943: 2 & 8; 1944: 40—Smith 1943: 283 & 289 (in part; records from “Tong-King (Chapa, Tam-dao” on p. 290); Deuve 1970: 84 & 88 (in part)—Tran et al. 1981: 380.
Amphiesma khasiensis (nec Tropidonotus khasiensis Boulenger, 1890 ).—Campden-Main 1970: 14.—Dao 1981: 7—Nguyen 1981: 10—Ho & Nguyen 1982: 139—Nguyen et al. 1995a: 32; 1995b: 96—Nguyen & Ho 1996: 68—Szyndlar & Nguyen 1996: 92—Orlov et al. 2000: 71 (in part)—Ziegler & Herrmann 2000: 53—Le et al. 2001: 27—Nguyen 2002: 37—Bain & Nguyen 2004: 7—Ho et al. 2004: 99—Hoang et al. 2005: 113—Nguyen et al. 2005: 74—Orlov 2005: 33.
Amphiesma khasiense (nec Tropidonotus khasiensis Boulenger, 1890 ).—Ziegler et al. 2006b: 262; 2007: 17, 22 (records of Tam Dao / Phong Nha), 28 & 37 (in part)—Nguyen et al. 2009: 355 (in part for the general distribition)—Das 2010: 332 (in part, only for the mention of northern Vietnam)—Bain & Hurley 2011: 104 & 129.
Amphiesma cf. khasiense .—Grismer et al 2007: 227—Nguyen et al. 2009: 629: Fig. 472 & 473.
Natrix khasiensis gilhodesi (nec Natrix gilhodesi Wall, 1925 , a junior subjective synonym of Tropidonotus khasiensis Boulenger, 1890 ).—Gée 1930: 76.
Natrix parallela parallela (nec Tropidonotus parallelus Boulenger, 1890 , a distinct species) –Bourret 1934b: 15 [Reprint: 5]; 1936b: 68 (in part; specimens from Cambodia).
Amphiesma inas (nec Natrix inas Laidlaw, 1901 , a distinct species).—Nabhitabhata et al. 2004: 124 (in part: mention of Khao Yai, Nakhon Ratchasima Province).
Material examined (95 specimens). People’s Republic of China. Yunnan Province. CIB 584172–584173, Pingbian Miaozu Zizhixian. Guizhou Province. CIB 63II 5098, Leishan. Guangxi Province. CIB 602339, Yao Shan; KFBG 9.VIII.1999, Jingxi. Jiangxi Province. MVZ 23622 (Paratype), “Hong San, alt. 850 meters, S. Kiangsi Province, south-eastern China (lat. 24° 58' N., long. 115° 50' E.)”, now near Jitan, Xunwu County. Guangdong Province. MVZ 23623 (Holotype), “Tai-yong, alt. 640 meters, E. Kwantung Province, south-eastern China (lat. 23° 34' N., long. 115° 55’ E), now Dayang, Jiexi County; SMNH 1238, no locality; ZMB 27694, “N-Kuangtung, China ”, i.e. northern Guangdong. Hainan Island and Province. CIB 745084, Diaoluo Shan. – Vietnam. Lao Cai Province. AMNH 153704, “Lao Cai”; MNHN 1935.0064, “Chapa”, now Sa Pa. Ha Giang Province. AMNH 148562, Yen Minh County. Lang Son Province. IEBR 2447, Mau Son. Vinh Phuc Province. IEBR 1290, MNHN 1935.0061–0063, MNHN 1935.0451–0453, MNHN 1958.0459–0460, MNHN 1997.3307, MNHN 1999.9093– 9094, MVZ 224141–224143, MVZ 224145–224148, MVZ 224153, MVZ 226513, MVZ 226515, PSGV 0 0 0 2 S, Tam Dao Hill Station. Quang Nam Province. AMNH 148552, Tra My; IEBR 2977, ROM 0 3848, ROM 21434, ROM 21438, Tay Gian. Kon Tum Province. IEBR 353, IEBR 1650–1652, IEBR 1654–1656, IEBR 1658–1662, IEBR 2537, IEBR 2540–2541. Gia Lai Province. FMNH 252117–252123, Ankhe. Lam Dong Province. BMNH 1921.4.1.3–4.1.5, BMNH 1969.1716–1718, Lang Bian Plateau, near Dalat; NCSM 77326, Bidoup-Nui Ba National Park, Lac Duong, 1,433 m, 12.16222°N, 108.665°E; NCSM 77328, NCSM 77333, Bidoup-Nui Ba National Park, Lac Duong, 1,494 m, 12.17528°N, 108.7006°E; NCSM 77334, Bidoup-Nui Ba National Park, Lac Duong, 1,484 m, 12.14417°N, 108.6772°E. No locality. CAS 73737, FMNH 71709, “South Vietnam ”; FMNH 178399, “ French Indochina ”. – Laos. Xiengkhuang Province. MNHN 1928.0056-0057, “Xiengkhouang”, former city of Xiengkhuang, now in the vicinity of Phonsavan. – Cambodia. Kampot Province. BMNH 1928.6.29.9, 1969.1710-1715, BMNH 1969.1720, “Bokor Mts.”, now Chuor Phnom Damrei, near Bokor Hill Station. Pursat Province. LSUHC 07442–07444, LSUHC 07464–07465, LSUHC 0 7484, Phnom Aural. – Thailand. Nakhon Ratchasima Province. FMNH 180153 – 180154, Khao Yai National Park.
Taxonomic comments. Amphiesma boulengeri has largely been identified in the literature as A. khasiense and A. modestum and, more rarely, A. inas and A. parallelum . These confusions can be traced in the chresonymy given above and are detailed in the Discussion below. It should be noted that the mentions of “ Natrix modesta modesta ” in Bourret (1935a: 259 & 261 [Reprint 1 & 3]), Bourret (1936b: 73–74 [in part], 74: Fig. 30) and Bourret (1937a: 29) refer to an undescribed species related to Amphiesma modestum , the description of which is in preparation. Both A. modestum and the undescribed species cannot be confused with A. boulengeri . The record of Tropidonotus modestus by Anderson (1879: 817) may also refer to another species related to A. modestum as currently defined but not to A. boulengeri (see also Pope 1935: 90).
Amphiesma boulengeri is here redefined as a wide ranging Indo-Chinese species. It is monotypic. Nevertheless, our results show noteworthy variations in the number of the ventral and subcaudal scales correlated with the geographical origin of specimens. It is possible to define a “northern” group, which includes populations present in China, northern and central Laos and northern Vietnam (provinces of Lao Cai, Ha Giang, Lang Son, and Vinh Phuc), a “southern” group, which contains populations from central and southern Vietnam (provinces of Quang Nam, Kon Tum, Gia Lai, and Lam Dong), and eastern Thailand, and a third group for populations of the Cardamom Mountains in south-western Cambodia. This variation is detailed below.
Diagnosis. A relatively large species of the genus Amphiesma characterized by the combination of (1) body slender in males, more robust in adult females; (2) nostrils laterally directed; (3) 25–28 maxillary teeth, gradually enlarged, the last 2 or 3 moderately enlarged; (4) 19 dorsal scale rows at midbody, weakly or moderately keeled, at the exception of scales of the 1st dorsal scale row which is entirely smooth; (5) dorsal pattern made of a distinct, cream or ochre (orange, coral, rusty red or reddish-brown in life) dorsolateral stripe on 5th–6th DSR with brighter dots or small blotches; (6) each posterior supralabial with a narrow, white, cream or ivory streak at mid height or on the lower part of the scale, often constricted near the edge of each supalabial and forming an irregular pale stripe starting behind eye or, more rarely, divided into elongate blotches; (7) pattern of the neck made of a distinct, narrow white streak extending from the corner of the mouth to the dorsolateral stripe; (8) venter ivory or cream, with the outer quarter of each ventral dark reddish-brown, separated from the dark dorsal colour by a narrow white line; (9) eye moderate to large, 1.4–2.6 times the distance between the lower margins of eye and of lip; (10) 136– 166 VEN, 83–119 SC; (11) internasals truncated anteriorly; (12) 9 (rarely 8 or 10) supralabials; and (13) 1 or 2 anterior temporals.
Variation (based on 95 examined specimens). Body cylindrical, elongate, rather slender in males, more robust in females, rather plump in old specimens; head elongate, moderately distinct from the thick neck, flattened; snout short, blunt, amounting for 23.5–29.6 % of HL or 1.2–1.9 times as long as diameter of eye; nostrils large, crescentic, directed laterally and piercing in the middle of the nasal; eye average, 1.5–2.5 times the distance between the lower margin of eye and the lower edge of lip, with a round pupil; tail long, thin and tapering.
The maximal total length recorded in the literature is 877 mm (Das 2010). The longest specimen examined by us is 770 mm long (SVL 524 mm; TaL 246 mm; BMNH 1969.1715; female). The longest known male is 649 mm (SVL 434 mm; TaL 215 mm; MVZ 23623; holotype). Ratio TaL / TL: 0.286–0.360, with a weak sexual dimorphism (see below).
Dentition. Maxillary teeth 25–28, gradually enlarged, the last 2 or 3 moderately enlarged.
Body scalation. DSR: 19–19–17 scale rows; dorsal scales rhomboedric, weakly to moderately keeled on DSR 2 to 9, normal in shape; scales of 1st DSR entirely smooth.
Scale row reductions: first reduction (19→17) at VEN 81–100 (x = 88.8; s = 2.3); second reduction (17→15) at VEN 86–103 (x = 92.2; s = 2.2).
VEN: 136–166 (plus 1 or 2 preventrals); SC: 83–115, all paired, without sexual dimorphism; ratio VEN / SC: 1.35–1.76; anal plate divided.
Position of the reduction to 6 scale rows around the tail: 5th–28th SC, with a strong sexual dimorphism. Ratio length of the portion of tail with 4 dorsal scale rows / length of the portion of tail with 6 dorsal scale rows: 1.03– 2.28 with a weak sexual dimorphism.
Our data show some variation in the number of ventrals and subcaudals related to the geographical origins of specimens. Three clusters can be defined as follows:
Cluster I: China + Northern Vietnam + Northern Laos + Central Laos
Cluster II: Central Vietnam + Southern Vietnam + Eastern Thailand
Cluster III: Cambodia (Cardamon Mountains)
Variation observed only in the numbers of ventrals, subcaudals and the two reductions of dorsal scale rows, are summarized in Table 3 View TABLE 3 .
Although populations referred to Clusters I and II differ mostly by the lower number of subcaudals in Cluster II, the populations from the Bokor and Cardamom mountain ranges, referred to Cluster III, differ clearly those of other clusters by all four characters retained here, especially the numbers of ventrals and subcaudals. Although these differences might be of taxonomic value, at present time we prefer to refrain from naming a new taxon for the population of the Cardamon Mountains as all other characters recorded on specimens of Cluster III are similar to those of other clusters.
Yang & Rao (2008: 255) cited two specimens with numbers of ventrals as high as 171 and 181 respectively. We consider these values well out of the range known for Chinese specimens and suspect a misidentification.
Head scalation. Rostral wider than high, only barely visible from above; nasals subrectangular, elongate, vertically divided on their lower half, with the posterior part larger and higher than anterior one; internasals subrectangular, in broad contact, about 0.8–1.2 times as long as wide, truncated but rather narrow anteriorly with anterior margin about 0.40–0.75 times the width of the posterior margin; 2 wide prefrontals, 1.1–1.6 (usually 1.2– 1.3) times as long as internasals; frontal shield-like with apex directed posteriorly, moderately enlarged, 1.1–1.7 (usually 1.3–1.4) longer than wide, 1.4–2.3 (usually 1.8–2.0) times as long as prefrontal; 1 supraocular on each side, longer than wide, about as wide as internasals; two wide parietals, variable in size, sometimes relatively short or usually elongate, 0.8–1.7 (usually 1.3–1.5) times longer than frontal; 1 / 1 subrectangular loreal scale, 0.6–1.0 times as high as long, in broad contact with the nasal; 9 (rarely 8, exceptionally 10) SL (9 / 9 in 79 cases out of 90, 8 / 8 in 4 cases, 8 / 9 in 6 cases, 9 / 10 in 1 case), 3rd–9th (or 8th or 10th) SL longer than high; 1st and 2nd SL small and short, in contact with nasal; 2nd and 3rd SL (exceptionally 2nd–4th SL) in contact with the loreal; 4th–6th SL entering orbit (exceptionally 4th–5th, 3rd–5th or 5th–7th SL, each in 1 case out of 90 specimens); 7th and 8th SL largest; 1–2 (exceptionally 3) preoculars on each side (2 preoculars in 56 / 180 occurrences; 1 preocular in 2 / 180 occurrences); 2–3 (exceptionally 1 in 1 / 180 occurrences) small postoculars; 1 anterior temporal (2 in 1 / 180 occurrences), narrow and elongate, with complete temporal formulas as 1 + 1 + 2, 1 + 2 + 2 or exceptionally 2 + 2 + 2; usually 10 / 10 infralabials (rarely 9 / 9, 9 / 10, exceptionally 10 / 11 or 11 / 11), first pair in contact with each with other, 1st– 5th IL in contact with anterior chin shields, 6th IL largest.
Coloration and pattern in alcohol. The dorsal surface of body is dark grey, dark grayish-brown, dark brown, chestnut brown or blackish-brown, distinctly darker, i.e. blackish-brown, on the five upper dorsal scale rows than on the sides; back with 2 rows and each side with 2 or 3 rows of small, blackish-brown spots, subrectangular or irregular, more or less well defined and scattered in alternance each with the other; on each side, a well-defined or even conspicuous, broad cream or ochre (orange, coral, rusty red or reddish-brown in life) dorsolateral stripe extends from the neck to the base of the tail on 5th and 6th dorsal scale rows, sometimes also on lower part of scales of the 7th row, usually visible throughout the body; on each dorsolateral stripe, often a series of large, elongate spots, brighter than the stripe, cream, yellow or ochre (coral, orange or red in life), more conspicuous and larger on the anterior part of the body, progressively fainter and vanishing into the stripe posteriorly, arranged every 2 to 4 dorsal; a short, narrow, ventrolateral stripe made of irregular cream spots extends from the neck to the anterior part of the body on the 1st dorsal scale row between the dark dorsal colour and the tips of ventrals, disappearing progressively after 10th–20th ventral; scales of the 1st dorsal scale rows sometimes paler in their center. The tail is as the body, with minute dorsolateral cream or yellow (coral, orange or red in life) spots in the extension of the dorsolateral stripes, vanishing progressively; scales of the 1st caudal rows paler in their center; posterior half of tail uniformly dark, usually variegated with irregular darker pigmentation.
The head is grayish-brown, brown, chestnut brown or dark brown, somewhat paler than the background colour of the body, often paler anteriorly above the snout than on the occiput, with irregular dark brown vermiculations or minute blackish-brown dots and with irregular paler areas on the frontal and parietal regions; sometimes a short cream streak just behind the suture of parietals; three or four anterior six supralabials largely white or cream (same in life), broadly edged with dark brown or black to such extent that these supralabials appear largely black; 5th and 6th supralabials more narrowly edged, appearing largely white and looking like two vertical streaks just below the eye; 7th to 9th supralabials with a conspicuously white, cream or ivory, more or less thick horizontal streak at mid height or on their lower part, this area forming either a broad, continuous stripe across the scales, or with elongate blotches, often constricted near the edge of each supalabial, in contact each with the other and forming an irregular but mostly continuous stripe, or, very rarely with separate, elongate blotches (see Fig. 5 View FIGURE 5 bottom); on each side of the neck, from the corner of the mouth behind 9th supralabial, an oblique, more or less thick but usually conspicuous, pure white or cream (same in life) oblique stripe (never isolate blotches), reaching the upper part of the neck; and connected to the dorsolateral stripe. The throat and chin are ivory or cream (pink in life); infralabials strongly and broadly edged with dark brown or blackish-brown; small, irregular dark brown spots on mentals present or reduced; sometimes minute dots on the preventral area. The pupil is black, the iris is dark brown in life.
The venter is uniformly ivory or cream (bright pink or reddish-gray in life); about a quarter of the outer part of each scale dark grey, dark brown or blackish-brown, separated from the dark similar colour covering the ventral tip by a narrow cream or ivory line crossing the scale at the base of the tip, producing a narrow, discontinuous ventrolateral stripe on each side of the venter and separating the dark colour of the tip from that of the inner part of the ventral; dark tips in contact without discontinuity with the background colour of the body except for a short distance on the anterior body, this pale space contributing the short ventrolateral stripe of the body. The tail is ivory or cream below (pink or red in life), with outer edges of subcaudals very dark brown; posteriorly, the tail becomes nearly entirely dark.
Hemipenis. In situ, it is short and stout, undivided but with two short extensions at its tip, reaching in situ the 6th or 7th SC; sulcus spermaticus simple, S-shaped, reaching the top of the hemipenis, with prominent lips; organ entirely covered with short but dense, thick spines, at the exception of the base which is covered with large calyces; lips of the sulcus smooth, spines not close to the sulcus; both extensions densely covered with rather numerous thin spines; 3 large, thick spines on the internal side of the organ at 90° of the sulcus; a very large spine at the base of the organ, on the side opposite to the sulcus.
Sexual dimorphism. It is weakly expressed in the ratio TaL/TL: males 0.294–0.360 (x = 0.334; s = 0.011); females: 0.286–0.340 (x = 0.317; s = 0.015) and (2) in the position (counted in subcaudals) of the reduction from 6 to 4 scale rows around the tail of subcaudals: males: 9th–28st SC (usually 15th–20th SC); females: 5th–23rd SC (usually 10–15 SC). We could not find any other dimorphism, especially not in the number of subcaudals (males: 84–119; females 83–115).
Distribution. As here defined, according to references cited in the chresonymy and our material, Amphiesma boulengeri is known from: People’s Republic of China. Recorded from the south and south-east of the country, in the provinces of Fujian, Jiangxi, Hunan, Guangdong (including Hong Kong), Hainan Island, Guangxi Autonomous Region, Guizhou, and Yunnan (east and centre of the province) (Zhao 2006).— Vietnam. This species is found throughout the country; it has been recorded from the provinces of Lào Cai (region of Sa Pa), Ha Giang, Cao Bang, Bac Thai, Bac Kan, Thai Nguyen, Vinh Phuc (Tam Dao), Ha Thay, Nghe An, Quang Binh (Phong Nha-Ke Bang National Parc), Quang Tri, Thua Thien-Hue (Bach Ma), Dak Lak, Kon Tum, and Lâm Dông (Da Lat) (after Nguyen et al. 2009). – Laos. Xiengkhuang Province. Currently known only from this northern province. — Cambodia. Recorded from the Cardamom Mountain, in the south-west of the country, in Kampot Province and Pursat Province.— Thailand. Definitely recorded from the province of Nakhon Ratchasima Province (Khao Yai National Park).
Biology. This species is a typical inhabitant of wet, primary evergreen submontane and montane forests and wet secondary forests between about 80 and at least 1,450 m a.s.l. but seems more common between 550 and 1,000 m a.s.l. Le et al. (2001) mentioned an elevation of 1,900 m for “ A. khasiense ” in Sa Pa Province, Vietnam. As A. khasiense , it is mostly found in damp areas along the edges of fast-flowing, rocky mountain streams, rice fields and on the forest floor. This snake is active both at day and at night. Terrestrial and semi-aquatic, A. boulengeri lives mainly among leaves of the forest litter and in low vegetation surrounding pieces of water; it readily enters shallow pieces of water such as forest creeks and puddles. Little else is known on its biology but it feeds on small frogs. It is oviparous. According to Gressitt (1941), the holotype contained three large, oblong eggs but nothing else is known on the reproductive habits of this species.
TABLE 3. Main variation in selected meristic characters in Amphiesma boulengeri.
| Sex / Cluster | Ventrals | Subcaudals | DSR Red. 1 | DSR Red. 2 |
|---|---|---|---|---|
| Males | ||||
| Cluster I | 140–152 x = 146.0 s = 3.1 | 93–109 x = 102.5 s = 3.9 | 85–97 x = 87.3 s = 3.8 | 85–99 x = 89.60 s = 3.8 |
| Cluster II | 136–159 x = 145.3 s = 6.1 | 84–101 x = 95.7 s = 4.3 | 81–91 x = 85.5 s = 4.1 | 83–97 x = 89.0 s = 4.8 |
| Cluster III | 155–165 x = 160.7 s = 3.1 | 108–119 x = 113.5 s = 4.5 | 86–100 x = 95.7 s = 4.6 | 87–103 x = 97.7 s = 5.2 |
| Females | ||||
| Cluster I | 139–151 x = 144.0 s = 3.4 | 90–100 x = 95.6 s = 3.6 | 82–89 x = 84.0 s = 3.4 | 83–90 x = 86.0 s = 2.7 |
| Cluster II | 136–153 x = 143.8 s = 5.6 | 83–101 x = 89.3 s = 4.7 | 82–97 x = 87.2 s = 5.5 | 86–103 x = 90.7 s = 4.7 |
| Cluster III | 152–166 x = 157.9 s = 5.7 | 99–115 x = 108.5 s = 5.7 | 92–100 x = 95.6 s = 5.7 | 98–103 x = 99.1 s = 5.7 |
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