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Hebius igneus DAVID, VOGEL, NGUYEN, ORLOV, PAUWELS, TEYNIÉ <br />& ZIEGLER, 2021

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Higher TaxaColubridae (Natricinae), Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Subspecies 
Common Names 
SynonymHebius igneus DAVID, VOGEL, NGUYEN, ORLOV, PAUWELS, TEYNIÉ
& ZIEGLER 2021
Tropidonotus modestus – BOURRET 1927 (?) (nec Tropidonotus modestus GÜNTHER 1875)
Natrix modesta – BOURRET 1936 (partim) (nec Tropidonotus modestus GÜNTHER 1875)
Natrix modesta – BOURRET 1939
Natrix modesta – BOURRET 1939
Natrix modesta – BOURRET 1939
Natrix modesta – TRAN et al. 1981 (?)
Amphiesma modesta – DAO 1981
Amphiesma modesta – DAO 1982: 8 (partim)
Amphiesma modesta – NGUYEN et al. 1995
Amphiesma modesta – ZHAO et al. 1998 (partim)
Amphiesma modesta – ORLOV et al. 2000
Amphiesma modesta – LE et al. 2001
Amphiesma modesta – BAIN & NGUYEN 2004
Amphiesma modesta – HO et al. 2005
Amphiesma modesta – NGUYEN et al. 2005 a
Amphiesma modesta – NGUYEN et al. 2005 b
Amphiesma modesta – ORLOV 2005 (partim)
Amphiesma modestum – ZIEGLER & LE 2006
Amphiesma modestum – NGUYEN et al. 2009 (partim)
Amphiesma modestum – ZIEGLER et al. 2007
Natrix modesta modesta – BOURRET 1935 (nec Tropidonotus modestus GÜNTHER 1875)
Natrix modesta modesta – BOURRET 1936 (partim)
Natrix modesta modesta – BOURRET 1937
Natrix modesta modesta – BOURRET 1939 (partim)
Amphiesma deschauenseei – NGUYEN et al. 2009 (partim) (nec Natrix deschauenseei TAYLOR 1934)
Amphiesma deschauenseei – BAIN & HURLEY 2011
Amphiesma deschauenseei – GUO et al. 2014 (partim) 
DistributionChina (Yunnan), Vietnam (Ha Giang, Lao Cai, Tuyen Quang; Vinh Phuc),Thailand (Nan)

Type locality: Ban Cai, 21°51’714N, 106°58’063E, elevation: 350–450m, Duc Xuan, Na Hang District, Tuyen Quang Province, Vietnam  
Reproduction 
TypesHolotype: IEBR 2295, adult female. Collected by Truong Quang Nguyen, Kien Van Doan and Cuc Thu Ho, 21 May 2004.
Paratypes (n = 8): IEBR, CIB, KIZ 
DiagnosisDiagnosis: A moderately to large sized species of the genus Hebius, defined by the combination of the following characters: (1) 19-19-17 dorsal scale rows, feebly or moderately keeled at midbody, more strongly keeled posteriorly at the exception of the scales of the 1st DSR, all smooth; (2) dorsal scales around the base of the tail strongly keeled; (3) head moderately distinct from the neck; (4) eye rather large; (5) maxillary teeth 29–30, the last two moderately enlarged; (6) tail long, with a ratio TaL/TL at least equal to 0.295; (7) VEN 159–169; (8) SC 115–129; (9) prefrontals 2; (10) one anterior temporal; (11) venter overall dark, i.e., dark brown, blackish-brown or black, with either pale irregular streaks or with several parallel, connected rows of dark blotches separated from each other by a narrow, irregular pale area; (12) dorsal surface and sides dark brown, dark chestnut-brown or blackish-brown, irregularly spotted by the presence of blackish-brown or black scales, darker than the background colour; (13) a dorsolateral series of bright orange, coral or rusty red blotches (yellow in preservative), distinctly enlarged behind the neck and at least on the anterior half of the body, then progressively smaller but always visible; these blotches may be connected and form a distinct dorsolateral line; (14) a dark postocular streak present; and (15) an ochre-yellow or yellowish-brown streak on each side of the neck and nape forming an incomplete collar (David et al. 2021).

Comparisons: Hebius igneus spec. nov. differs from H. annamensis (Bourret), H. atemporalis (Bourret), H. chapaensis (Bourret), H. groundwateri (Smith), H. nigriventer (Wall) new. comb., H. sauteri (Boulenger) (including the subspecies H. s. bourreti and H. s. maximus), H. taronensis (Smith), and H. venningi (Wall), as well as three species present in Borneo, i.e., H. arquus (David & Vogel), H. frenatus (Dunn) and H. sarawacensis (Günther), in having 19 DSR vs. 17 rows in all these species and sometimes 15 in H. annamensis. Furthermore, by its largely black venter, Hebius igneus spec. nov. further differs from the mainland species H. annamensis, H. atemporalis, H. groundwateri, H. sauteri, and H. venningi which have a large part of the venter pale.
Among the numerous species with 19 dorsal scale rows, Hebius igneus spec. nov. differs by its dorsal pattern from a number of distinctly striped species such as Amphiesma stolatum (Linnaeus), and Hebius bitaeniatus (Wall), H. clerki (Wall), H. metusia (Inger, Zhao, Shaffer & Wu), H. octolineatus (Boulenger) and H. parallelus (Boulenger) which display pale and dark dorsal stripes. Furthermore, these species have a pale venter and a lower number of subcaudals.
Also by its dorsal and ventral patterns, Hebius igneus spec. nov. differs from species with 19 dorsal scale rows of the Hebius khasiensis-group, hence containing H. khasiensis (Boulenger), H. boulengeri (Gressitt), H. craspedogaster (Boulenger), H. inas (Laidlaw), H. johannis (Boulenger), H. kerinciensis (David & Das), and H. leucomystax (David, Bain, Nguyen, Orlov, Vogel, Vu & Ziegler). All these species have dorsolateral spots and stripes but the blotches are smaller and all these species have a pale venter at the exception of the tips of ventrals. Furthermore, both H. khasiensis and H. inas differ from Hebius igneus spec. nov. by the pattern of their supralabials, composed of distinct, pale blotches in H. khasiensis and H. inas, at least on the posterior supralabials. Furthermore, Hebius igneus spec. nov. differs from H. leucomystax (David, Bain, Nguyen, Orlov, Vogel, Vu & Ziegler) and H. lacrima Purkayastha & David by its overall dark supralabials vs. supralabials largely white in these two latter species, plus by its venter largely dark, and its different dorsal pattern.
In the same way, Hebius igneus spec. nov. differs from H. optatus (Hu & Zhao) and H. andreae (Ziegler & Le) by its dorsal pattern made of conspicuous dorsolateral blotches vs. patterns made of vertical, narrow or wide crossbars, without longitudinal dorsolateral stripes or series of dots in these two latter species. Malnate (1962) referred Hebius craspedogaster, H. popei, H. pryeri, H. sauteri and H. vibakari in a distinct species group, named as the “Amphiesma vibakari-group”. Besides H. sauteri, which has 17 DSR at midbody, Hebius igneus spec. nov. differs from these species by its dorsal and ventral patterns and by its higher number of subcaudals. Hebius igneus spec. nov. obviously shares one or more characters with other species of “dark-bellied” Hebius, as treated here. However, for summarizing, Hebius igneus spec. nov. differs from H. annamensis, H. chapaensis, H. deschauenseei, H. modestus, H. nigriventer, H. taronensis and H. venningi by the combination of the following characters: (1) 19 DSR at midbody, (2) scales near vent and the base of the tail strongly keeled, (3) at least 159 ventrals, (4) a dorsolateral series of bright orange, coral or rusty red elongate blotches, larger anteriorly but extending behind half of the body, and (5) several parallel series of dark blotches on the venter more or less separated by narrow pale stripes. Tentatively included in this group but not addressed here, Hebius xenura (Wall), which has also 19 DSR at midbody and dorsal blotches, differs from Hebius igneus spec. nov. by (1) its undivided subcaudals (vs. divided in H. igneus), (2) fewer subcaudals, i.e., 92–105 vs. 115–129, and (3) its ventral pattern, pale with tips of ventrals dark vs. venter overally dark.
Hebius annamensis, H. chapaensis, H. nigriventer, H. venningi, and H. taronensis have 17 DSR at midbody (or 15 in H. annamensis) vs. 19 in Hebius igneus spec. nov. Furthermore, H. annamensis and H. venningi have a venter largely pale mesially or, in H. venningi, pale mesially on the anterior part of the body, clouded with darker hues of brown on the outer parts of ventrals and entirely clouded posteriorly. Among the other two species of this group with 19 DSR at midbody, Hebius igneus spec. nov. differs from H. modestus by (1) dorsal scales before the base of the tail and supracaudal scales just after strongly keeled vs. weakly keeled in H. modestus, (2) slightly more ventrals, 159–169 VEN vs. 143–163, (3) venter overall dark brown, blackish-brown or black, with either pale irregular streaks or with several parallel, connected rows of dark blotches separated from each other by a narrow, irregular pale area vs. venter pale ochre-brown or pale brown on a wide central area, broadly edged with dark brown or blackish-brown on the outer quarter of ventrals, rarely nearly overally dark except on its anterior part, (4) a dorsolateral series of bright orange, coral or rusty red blotches, distinctly enlarged behind the neck and at least on the anterior half of the body, then progressively smaller but always visible vs. a more or less conspicuous, ochre-yellow, ochre-red, orange-brown or reddish-brown stripe, often reduced to a succession of elongate blotches on the anterior part of the body, and (5) a dark postocular streak present vs. no such streak.
Lastly, Hebius igneus spec. nov. is more similar to H. deschauenseei, from which it differs as follows: (1) supracaudal scales around the base of the tail strongly keeled vs. weakly keeled in H. deschauenseei, (2) Ratio TaL/TL, 0.30–0.33 vs. 0.33–0.38, (3) venter overally dark after the 5th–8th ventrals, i.e., dark brown, blackish-brown or black, with 4 or 5 parallel, connected rows of dark blotches separated from each other by a narrow, irregular pale area vs. venter never entirely dark along its whole length, creamish-yellow or pale yellowish-grey with three parallel stripes of aligned dark blotches separated from each other by a pale area in H. deschauenseei, (4) dorsal surface and sides dark brown, dark chestnut-brown or blackish-brown vs. olive-brown, olive-grey or greyish-brown, (5) dorsolateral blotches, distinctly enlarged behind the neck and at least on the anterior half of the body, then progressively smaller but always distinct vs. dorsolateral blotches not as enlarged anteriorly and progressively smaller and united, forming a dorsolateral stripe, and (6) a dark postocular streak present vs. postocular streak absent or barely visible (DAVID et al. 2021).

Color in life: Dorsal surface and sides chestnut-brown, dark brown, blackish-brown or even deep black, more or less iridescent, nearly uniform or irregularly spotted by the presence of blackish-brown or black scales, darker than the background colour; in some specimens, many scales of the sides are marked with yellow or yellowish-ochre; a series of 65 to 75 conspicuous, elongate dorsolateral blotches on 5th and 6th or 6th and 7th dorsal scale rows, bright yellow, orange, rusty red or bright reddish-ochre, distinctly enlarged on the neck where they are about 2 to 2.5 DSR long and 1.5 to 2 scales high, then progressively smaller, often split into two parts on fore third of the body, irregularly split after midbody and reduced to a spot or a dash posteriorly but always visible; dorsolateral stripe usually absent but a faint, ochre-brown or reddish-brown stripe, may be present between the blotches along the posterior part of the body. The upper surface of tail is dark colored as the body, either with a dorsolateral series of horizontally elongate pale spots in preservative or uniformly dark.
Head: upper surface, parietal region and background colour of supralabials dark brown or blackish-brown, with scattered bright yellow, golden yellow or yellowish-ochre beige or pale brown dots on internasals, prefrontals and cephalic region; a short, cream sagittal line just behind the suture between parietals present but more or less faint; rostral and sides of the snout slightly paler than upper head surface; a yellowish-brown or yellowish-ochre line on the outer edge of each parietal and onto upper temporals very thin or often absent; usually (rarely completely absent) a yellowish-brown or yellowish-ochre postocular streak, varying from barely visible or even reduced to a few dots, to quite broad and distinct, extending from upper postocular downwards to 9th SL and the corner of the mouth; 1st to 6th supralabials more or less distinctly paler, i.e., dark brown or dark ochre-brown, near their anterior edge, also often in their central part or even the anterior half of the scale; 6th SL usually more distinctly paler in its anterior half; 7th SL yellowish-cream or ochre-brown on a large part, either as a triangular pale area, wider at its base on its anterior half or on its central area with only the top and bottom of the scale blackish-brown; 8th SL also yellowish-cream or ochre-brown on its lower anterior part, sometimes forming with the central area of 7th SL a conspicuous, elongate and oblique marking, or when the paler areas of the 7th and 8th are disjunct, forming two pale, parallel triangles; posterior part of 9th SL creamish-yellow or pale yellowish-brown; upper parts of 7th, 8th and anterior part or middle of 9th SL, and anterior temporal uniformly black or very dark blackish-brown, forming a large, irregular, oblique postocular streak extending from behind the eye to the corner of the mouth and slightly beyond in some specimens; sides of the neck yellowish-cream, strongly spotted with blackish-brown, from which extends a yellowish-cream, more or less wide oblique streak directed upwards and backwards, forming an irregular, yellowish-brown V-like mark just behind the head with its branches not in contact on the neck.
Venter overall dark with numerous dark greyish-brown, dark brown or blackish-brown blotches on a yellowish-cream background colour; venter largely yellowish-cream anteriorly on the first 5 to 8 ventrals, with a mesial row of dark blotches or a solid streak and tips of ventrals dark, then, from 3 to 5 rows of parallel, elongate dark blotches, one to three mesially plus one on the tip of each ventral, separated by pale background colour, extending from the anterior margin of the ventral and not or barely reaching its posterior margin, these blotches becoming progressively wider and connected, reducing the background colour to short, pale irregular streaks separating the rows of dark blotches; after the first third to half of the body, depending on specimens, the blotches are entirely united and form dark, transversal blotches on much of each ventral, leaving a narrow, pale area along the posterior edge of each ventral. Tail blackish-brown below, with only the posterior edge of each subcaudal narrowly edged with beige brown or yellowish-ochre. 
CommentDiet: Frogs of the families Megophryidae and Ranidae (DAVID et al. 2021) 
EtymologyThe specific nomen is the Latin adjective igneus (a, um) which means “in fire”. This specific name was coined by analogy to the large bright orange dorsolateral blotches on the forepart of the body. 
References
  • BAIN, R.H. & NGUYEN, T.Q. 2004. Herpetofaunal diversity of Ha Giang Province in Northeastern Vietnam, with descriptions of two new species. American Museum Novitates, 3453: 1–42
  • Bourret, R. 1936. Les Serpents de l'Indochine, vol. 2. Imprimerie Henri Basuyau & Cie, Toulouse.
  • BOURRET, R. 1937. Notes herpétologiques sur l’Indochine française. XIII. Serpents récemment récoltés au Tonkin et en Annam. Bulletin général de l’Instruction Publique, 16e Année (1936–1937), 9 Mai 1937, Annexe, 27–36
  • BOURRET, R. 1939. Notes herpétologiques sur l’Indochine française. XX. Liste des Reptiles et Batraciens actuellement connus en Indochine Française. Bulletin général de l’Instruction Publique, 19e Année (1939–1940), 4 Décembre 1939, Annexe, 49–60
  • BOURRET, R. 1939. Notes herpétologiques sur l’Indochine française. XIX. La faune herpétologique des stations d’altitude du Tonkin. Bulletin général de l’Instruction Publique, 19e Année (1939–1940), 4 Décembre 1939, Annexe, 41–47
  • Bourret,R. 1935. Notes herpétologiques sur l'Indochine Française X. Les serpents de la station d’altitude du Tam-dao. Bull. Gen. Instr. Pub. Hanoi 8 (avril): 259-271 (1-13)
  • Dao Van Tien 1981. [List of the Vietnamese snakes at present known in Vietnam]. Tap Chi Sinh Vat Hoc 3(1): 6-10.
  • DAO, T.V. 1982. Key to the species of Vietnamese snakes. Part II. Tap chi Sinh Vat Hoc, 4 (1), 5–9. [in Vietnamese]
  • DAVID, PATRICK; GERNOT VOGEL, TRUONG QUANG NGUYEN, NIKOLAI L. ORLOV, OLIVIER S. G. PAUWELS, ALEXANDRE TEYNIÉ, THOMAS ZIEGLER 2021. A revision of the dark-bellied, stream-dwelling snakes of the genus Hebius (Reptilia: Squamata: Natricidae) with the description of a new species from China, Vietnam and Thailand. Zootaxa 4911: 1–61 - get paper here
  • GUO, PENG; FEI ZHU, QIN LIU, LIANG ZHANG, JIAN X. LI, YU Y. HUANG & R. ALEXANDER PYRON 2014. A taxonomic revision of the Asian keelback snakes, genus Amphiesma (Serpentes: Colubridae: Natricinae), with description of a new species. Zootaxa 3873 (4): 425–440 - get paper here
  • NGUYEN, S.V., NGUYEN, T.Q. & HO, C.T. 2005. ) Species composition of Amphibians and Reptiles from Vanban District, Laocai Province. [in Vietnamese, English summary] Tap Chi Sinh Hoc, 27 (4A), 117–123.
  • Orlov, N. L., R. W. Murphy & T. J. Papenfuss 2000. List of snakes of Tam-Dao mountain ridge (Tonkin, Vietnam). Russ. J. Herpetol. 7 (1): 69-80. - get paper here
  • Ziegler, T., and Quyet, L.K. 2006. A new natricine snake of the genus Amphiesma (Squamata: Colubridae: Natricinae) from the central Truong Son, Vietnam. Zootaxa 1225:39-56 - get paper here
  • ZIEGLER, THOMAS; RALF HENDRIX, VU NGOC THANH, MARTINA VOGT, BERNHARD FORSTER & DANG NGOC KIEN 2007. The diversity of a snake community in a karst forest ecosystem in the central Truong Son, Vietnam, with an identification key. Zootaxa 1493: 1-40 - get paper here
 
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