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Accepted by Z. Nagy: 9 Dec. 2014; published: 17 Feb. 2015
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2015 Magnolia Press
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http://dx.doi.org/10.11646/zootaxa.3919.2.9
http://zoobank.org/urn:lsid:zoobank.org:pub:B9AA0822-2908-4C4C-A14A-30007F40F9BE
Revalidation of Natrix clerki Wall, 1925, an overlooked species in the genus
Amphiesma Duméril, Bibron & Duméril, 1854 (Squamata: Natricidae)
PATRICK DAVID
1
, ISHAN AGARWAL
2
, RAMANA ATHREYA
3
, ROSAMMA MATHEW
4
,
GERNOT VOGEL
5
& VIRAL K. MISTRY
6
1
Reptiles & Amphibiens, UMR 7205 OSEB, Département Systématique et Évolution, CP 30, Muséum National d’Histoire Naturelle, 57
rue Cuvier, F-75231 Paris Cedex 05, France. E-mail: pdavid@mnhn.fr
2
Centre for Ecological Sciences, Indian Institute of Science, Bangalore 560012, India. E-mail: ishan.agarwal@gmail.com
3
Indian Institute of Science Education & Research, Pune 411008, Maharashtra, India. E-mail: rathreya@iiserpune.ac.in
4
B-4, 502, Rajyog Township, Sinhgad Road, Wadgaon Kh., Pune-411 041, India. E-mail: rosamma@rediffmail.com
5
Society for Southeast Asian Herpetology, Im Sand 3, D-69115 Heidelberg, Germany. E-mail: Gernot.Vogel@t-online.de
6
9B/305-306, Whispering palms, Kandivali east, Mumbai 400101, Maharashtra, India. E-mail: mistryviral@gmail.com
Abstract
Natrix clerki Wall, 1925, previously known from its sole holotype and considered a synonym of Amphiesma parallelum
(Boulenger, 1890), is resurrected in the genus Amphiesma on the basis of the analysis of morphological variation in 28
specimens of “Amphiesma parallelum” auctorum, plus six living, unvouchered specimens discovered in Arunachal
Pradesh and Nagaland, India, and one vouchered specimen from Talle Valley in Arunachal Pradesh. Specimens from
northeast India (Nagaland), northern Myanmar, and China (Yunnan), previously identified as Amphiesma parallelum ei-
ther in the literature or in museum’s catalogues, are also here referred to A. clerki. The holotype of Amphiesma clerki is
redescribed. As a consequence, the definition of Amphiesma parallelum is modified. A. parallelum inhabits the Khasi
Hills and Naga Hills in Northeast India, whereas A. clerki has a wider range in the Eastern Himalayas, northern Myanmar
and Yunnan (China). Amphiesma clerki differs from A. parallelum by its longer tail, dorsal scales more strongly keeled,
scales of the first dorsal scale row strongly keeled vs. smooth, a postocular streak not interrupted at the level of the neck,
and a much more vivid pattern on a darker background colour. Characters of species of the Amphiesma parallelum group,
i.e. A. clerki, A. parallelum, A. bitaeniatum, A. platyceps and A. sieboldii are compared. A key to this group is provided.
Key words: Reptilia, Nepal, India, Myanmar, China, taxonomy
Introduction
The taxonomy of the Asian natricid genus Amphiesma Duméril, Bibron & Duméril, 1854 remains in a state of flux,
with several species described or resurrected during the last 15 years (David & Das 2003; David et al. 2005, 2007,
2010, 2013; Ziegler & Quyet 2006). Currently, this large genus is composed of 42 species. Malnate (1960)
addressed the systematics of this genus, previously considered a synonym of the large genus Natrix Laurenti, 1768.
Malnate defined a group named as “Amphiesma khasiense species group” that included A. khasiense (Boulenger,
1890), A. craspedogaster (Boulenger, 1899), A. popei (Schmidt, 1925), A. pryeri (Boulenger, 1887), A. sauteri
(Boulenger, 1909), A. vibakari (Boie, 1826), A. venningi (Wall, 1910) as well as A. parallelum (Boulenger, 1890)
and related species. However, Malnate (1960) had pointed out that Amphiesma parallelum and the related, striped
species differed from other members of his “A. khasiense group” by their abruptly and strongly enlarged posterior
maxillary teeth. On the basis of these characters, David et al. (2013) referred Amphiesma parallelum and related
species into a distinct, informal species group, that currently includes Amphiesma parallelum (Boulenger, 1890), A.
bitaeniatum (Wall, 1925), A. octolineatum (Boulenger, 1904), A. platyceps (Blyth, 1854), and A. sieboldii
(Günther, 1860). Furthermore, A. metusia Inger, Zhao, Shaffer & Wu, 1990, a striped species, is tentatively referred
to this group due to its color pattern. Species of the A. parallelum group, that inhabit the Himalayas and
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neighbouring mountain ranges, are all characterized by a more or less striped pattern and are morphologically
rather similar. Unsurprisingly, the confusion within this species group has long been extensive in the literature. For
example, Amphiesma platyceps and A. sieboldii have been confused with A. parallelum, whereas A. bitaeniatum
has also been largely confused with A. parallelum, A. octolineatum and A. metusia (see, for example, Bourret 1936;
Smith 1943).
Smith (1943) adopted a conservative approach in the systematics of Amphiesma parallelum. Without
comment, he placed in its synonymy A. bitaeniatum, a taxon recognized by Pope (1935) in spite of consistent
differences with A. parallelum (see David et al. 2005). Smith (1943) also placed in the synonymy of this latter
species Natrix clerki, a taxon described by Wall (1925: 809) from a single specimen (BMNH 1946.1.13.50),
obtained in “Sinlum Kaba, Kachin Hills”, now Sinlumkaba, Kachin State, Myanmar. In the frame of various
investigations on the genus Amphiesma, the holotype of Natrix clerki, was examined by PD. It was briefly
mentioned in David et al. (2005) who suggested that Natrix clerki Wall, 1925 might prove to be a valid species,
along with two slightly different specimens identified as Amphiesma parallelum by David et al. (2005).
Recent herpetological surveys in Northeast India (Ashok Captain, pers comm; Agarwal et al. 2010) resulted in
the discovery of five additional specimens, four in Arunachal Pradesh and one in Nagaland, sharing the diagnostic
features of the holotype of Natrix clerki. This newly collected material and the examination of the holotype of
Natrix clerki led us to investigate in details the variation in A. paralellum auctorum. This analysis allowed the
identification of a number of morphological differences between these specimens and those morphologically
similar to the lectotype of Amphiesma parellum. Morphological variation in “Amphiesma parallelum” is discussed
in detail and two species are recognized.
Material and methods
This study is based on the examination of 138 preserved specimens, including the name bearing types of Natrix
clerki Wall, 1925 and Tropidonotus parallelus Boulenger, 1890, plus six unpreserved specimens, observed alive,
from three localities in Arunachal Pradesh, India, and one specimen from Nagaland (see Agarwal et al. 2010). Four
of the specimens from Arunachal Pradesh were examined in detail and photographed but were not preserved in
agreement with Indian regulations. The living specimens are as follows: Arunachal Pradesh (AP): specimens AP
1–2, Eaglenest Wildlife Sanctuary, (Bompu camp), Dafla Hills, West Kameng District; Specimens AP 3–4,
outskirts of Shidi (Gandhigram), Changlang District; Specimen RS09 from Talle Valley, now deposited in IISER,
Pune. Nagaland: no number, near the Tragopan Sanctuary (25.63549N-094.01261E), Khonoma. Specimens of
Amphiesma clerki and A. parallelum examined are listed under the species accounts. Specimens of other species
referred to the Amphiesma parallelum group are listed in the Appendix.
All measures are in millimetres. Body and tail lengths were measured to the nearest millimetre in preserved
specimens. Measurements of living snakes were taken to the nearest centimetre. Ventral scales on preserved
specimens were counted according to Dowling (1951). Ventral counts on living, unpreserved specimens AP1–4,
began at the first discernible ventral, considered to be the of the largest ventral scale, at least twice as large as the
preceding scales, without regard for the point of origin of the dorsal scale rows. Our counting method for living
specimens resulted in counts that are approximately two scales greater than Dowling’s method on preserved
specimens. The terminal scute was excluded from counts of the number of subcaudals. The number of dorsal scale
rows is given at one head length behind head, at midbody (i.e. at the level of the ventral plate corresponding to half
of the total ventral number), and at one head length before vent, respectively. Values for symmetric head characters
are given in left/right order. Measurements made on live snakes are likely to not have the same level of precision as
preserved material, but we present these data as valuable information for this poorly known taxon.
Abbreviations used for measurements and meristic characters are:
Measurements and proportion. HL: head length; SVL: snout-vent length; TaL: tail length; TL: total length;
TaL/TL: proportion tail length / total length.
Meristic characters: ATe: anterior temporals; DSR: dorsal scale rows; K1SR : keeling of first DSR at
midbody; KSR: keeling of dorsal scale rows at midbody; NSR: notching of apical part of dorsal scales; PreOc:
preocular(s); SC: subcaudals; SL: supralabials; VEN: ventrals.
Museum abbreviations. AMNH—The American Museum of Natural History, New York, USA. BMNH—The
Natural History Museum, London, United Kingdom. CAS—California Academy of Sciences, San Francisco, USA.
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REVALIDATION OF NATRIX CLERKI WALL, 1925
CTNRC—Center for Thai National Reference Collections, National Research Council of Thailand, Bangkok,
Thailand. FMNH—The Field Museum, Chicago, USA. IISER—Indian Institute of Science Education and
Research, Pune, India. KSC—Kohima Science College, Kohima, India. MHNG—Museum d’Histoire Naturelle,
Geneva, Switzerland. MNHN—Muséum National d’Histoire Naturelle, Paris, France. NMW—Naturhistorisches
Museum Wien, Vienna, Austria. ROM—Royal Ontario Musem, Toronto, Canada. SMNH—Shanghai Museum of
Natural History, Shanghai, People’s Republic of China. THNHM—Thailand Natural History Museum, Pathum
Thani, Thailand. USNM—United States National Museum, Washington, USA. ZMB—Zoologisches Museum für
Naturkunde der Humboldt-Universität zu Berlin, Berlin, Germany. ZSI—Zoological Survey of India, Kolkata,
India. ZSI/ERS—Zoological Survey of India (Eastern Regional Station), Shillong, India.
Results
On the basis of our data from preserved specimens, supplemented by the fsix living specimens, we can distinguish
two distinct forms: Form 1 is characterized by (1) a dorsal pattern made of two broad, conspicuous dorsolateral
stripes on a rather pale (in preservative), uniform background, (2) a postocular streak not in contact on the side of
the neck with the dark lateral stripe, (3) scales of the first dorsal row not keeled, other scale rows weakly keeled, (4)
a comparatively short tail, and (5) posterior maxillary teeth moderately enlarged, always separated from other teeth
by a distinct diastema. Form 2 is characterized by (1) a strongly distinct but confused pattern, made of a mixture of
pale blotches, dark spots and faint dorsolateral stripes on a dark background, (2) a narrow postocular streak
continuous on neck, (3) a comparatively long tail, (4) all dorsal scale rows strongly keeled, and (5) posterior
maxillary teeth strongly enlarged, not separated from other teeth or only by a short diastema. These results are
summarized in Tables 1–2.
Comparison of the type specimens of Tropidonotus parallelus Boulenger, 1890 and Natrix clerki Wall, 1925
leads us to refer all specimens of Form 1 to Amphiesma parallelum, whereas all specimens of Form 2, as well as
five living Indian specimens, agree very well with the holotype of Natrix clerki. As no intermediate condition has
been found and as these differences are correlated with the geographic distribution, we argue that Natrix clerki
Wall, 1925 is distinct from Amphiesma parallelum. Therefore, Natrix clerki Wall, 1925 is resurrected from the
synonymy of A. parallelum. The holotype of Natrix clerki is described in detail, and variation and distribution of
this species are presented herein on the basis of data from the 10 preserved and five photographed specimens
available to us.
Amphiesma clerki (Wall, 1925)
(Fig. 1–6)
Natrix clerki Wall, 1925: 809. Type locality. “Sinlum Kaba, Kachin Hills”, now Sinlumkaba (24°16'N–97°31'E), Kachin
State, Myanmar. Holotype. BMNH 1946.1.13.50, adult male; collected by Mr. Clerk, 1924.
Material (n = 10 preserved specimens + 5 unpreserved, living specimens). Nepal (?). BMNH 58.6.24.5,
“Nepal”.—India. Sikkim
. BMNH 60.3.19.1359, NMW 22383:2, “Sikkim”, no precise locality.—West Bengal.
BMNH 80.11.10.153, “Darjeeling”; BMNH 1923.10.13.38, “Darjeeling District”.—Arunachal Pradesh
. IISER
RS09, Talle Valley.— Nagaland
. KSC 414, Sechu, 1000 m. —Myanmar. Kachin State. BMNH 1946.1.13.50
(Holotype), “Sinlum Kaba, Kachin Hills”, now Sinlumkaba; BMNH 1940.6.4.29, “Pangnamdim, The Triangle,
Upper Burma”, a village about 24 km northeast of Watamkawng (27°43'N-97°52'E).—People’s Republic of
China. Yunnan Province
. CAS 215036, Nu Jiang Nature Reserve, near Pianma (26°00'10.3 N-98°39'31.7 E), Nu
Jiang County.
Systematics. This species has seemingly been mentioned in the literature only by Wall (1925, 1926).
Subsequently, it was placed in the synonymy of Natrix parallela (now Amphiesma parallelum) by Smith (1943:
288). It is here resurrected with full species status. This species is monotypic.
Diagnosis. A species of Amphiesma characterized by (1) a dark dorsal background, dark brown, dark greyish-
brown or dark brownish-grey (same in life), becoming more or less distinctly darker on the back than on sides; (2)
laterally, a series of pale yellow, pale yellowish-brown or brownish-yellow spots (reddish-brown or rusty red in
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life), distinctly larger and conspicuous on the neck and forepart of the body, connected along the hinder half of the
body by a pale dorsolateral stripe extending on the 5
th
–7
th
dorsal scale rows; (3) scales broadly edged with dark
brown on the lower part of the sides, producing irregular blotches; (4) faint, irregular and short blackish-brown
streaks on the 1
st
dorsal scale row, producing a very discontinuous zigzag ventrolateral stripe; (5) a conspicuous,
broad, blackish-brown postocular streak extending from behind the eye to the corner of the mouth then extending,
usually without gap (rarely discontinuous) on the side of the neck; (6) a conspicuous, pale (cream or pale yellow),
V or Y-shaped chevron starting on each side from behind the last supralabial and reaching the upper surface of the
neck, pointing backwards; (7) supralabials dotted with brown to black dots or even nearly brown, edged with dark
brown and with usually a conspicuous streak on the hinder margin of the 6
th
SL; (8) venter creamy yellow (bright
reddish-brown, rusty red or crimson in life), with outer tips of ventral scales light brown (dark purplish-red in life);
(9) 18–22 maxillary teeth, the last two strongly enlarged, either without diastema or barely separated from anterior
teeth by a short diastema; (10) 19–19–17 dorsal scale rows; (11) dorsal scales strongly keeled on all dorsal scale
rows, including 1
st
row; (12) tail long, ratio tail length / total length 0.262–0.325 (0.280–0.325 in males,
0.262–0.273 in females); (13) 162–173 ventrals; (14) 85–108 subcaudals; (15) anal plate divided; (16) usually 1
preocular; (17) usually 2 anterior temporals.
Redescription of the holotype of Natrix clerki, BMNH 1946.1.13.50 (Fig. 1–2)
Habitus. Body elongate, cylindrical; head elongate, rather narrow, distinct from the neck; snout average,
accounting for 26.3 % of HL, 1.3 times as long as eye diameter, slightly flattened, narrowing anteriorly, blunt when
seen from above, rounded seen in profile, with no defined canthus rostralis; nostril piercing laterally; eye large, its
diameter 2.2 times the distance between its inferior margin and upper lip edge; pupil round; tail long, cylindrical
and progressively tapering.
SVL 384 mm; TaL 170 mm; TL 554 mm. Ratio TaL / TL: 0.307.
Dentition. Maxillary teeth: 21 subequal + 3 strongly and abruptly enlarged teeth, about twice as large as
anterior teeth, without diastema.
Body scalation. Dorsal scales: 19-19-17 rows; scales very strongly keeled on the upper part of the back, a little
bit less strongly keeled on the sides and the 1
st
dorsal scale rows; all scales strongly and deeply notched at their
apical part, especially on the posterior part of the body; apical pit present on each scale; scales of 1
st
dorsal scale
row slightly enlarged.
Ventrals: 171 (+ 2 preventrals); subcaudals: 108, all paired; anal divided.
Head scalation. Rostral 1.6 times as wide as high; nasal rectangular, longer than high, with a lateral nostril
piercing in the middle of the scale; internasals subtriangular, slightly (1.1 times) longer than wide, anteriorly
narrowed but truncated, with the fore width about 0.6 times as great as the hind width; prefrontals barely longer
than internasals; one large, longer than wide undivided supraocular on each side; frontal bell-shaped, 1.7 times as
long as wide; parietals large, in contact along a suture about 0.9 times as long as frontal; one subrectangular loreal
on each side, 1.1 times longer than high; 8 / 8 supralabials, 1
st
and 2
nd
supralabials in contact with nasal, 2
nd
and 3
rd
supralabials in contact with loreal, 3
rd
–5
th
supralabials entering orbit, 7
th
supralabial largest; 1 / 1 tall, narrow
preocular; no subocular; 3 / 3 small postoculars; 1 anterior temporal, temporals with the formula 1+2 on each side;
9 / 9 infralabials, 1
st
pair in contact, 1
st
–4
th
infralabials in contact with anterior chin shields, 4
th
and 5
th
infralabials
largest.
Coloration and pattern. Body dark greyish-brown on the back, brown or tan on its sides but with a distinctly
darker, blackish-brown area on the neck sides and extreme forepart of the body; on the sides of the body, most
scales of the 1
st
to 4
th
dorsal scale rows broadly edged with blackish-brown, producing reticulations and blackish-
brown, irregular, oblique blotches on the sides; dorsum irregularly marked with small, black spots more or less
arranged as one or two vertebral series and two irregular series on each outer side of the dorsum just above each
dorsolateral stripe; on each side, a series of large, rounded, irregular but conspicuous pale yellow blotches on the
neck and fore part of body, extending on 4
th
–7
th
or 5
th
–7
th
dorsal scale rows, not connected by a dorsolateral stripe,
progressively turning into smaller, pale yellowish-brown blotches aligned on the upper half of 5
th
, 6
th
and lower half
of 7
th
DSR, rounded on the fore quarter to third of the body, more elongate, faint and reduced to the 5
th
and 6
th
rows
posteriorly, connected from the first quarter to third of the body by an indistinct pale yellowish-brown dorsolateral
stripe extending along on the 5
th
DSR from the second quarter of the body to the vent, more conspicuous
posteriorly; another series of 7 or 8 vertical, yellow streaks interrupting the dark brown dorsal colour of the sides of
the first quarter of the body, originating from the ventral colour and reaching the levels of the 5
th
or 6
th
DSR,
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REVALIDATION OF NATRIX CLERKI WALL, 1925
alternating or in contact with the dorsolateral blotches; scales of the 1
st
DSR marked with faint and irregular
blackish-brown short streaks, producing a very discontinuous, zigzag-like ventrolateral stripe, more conspicuous
from the level of the second quarter of the body. Upper tail surface as the body, with scales strongly edged with
dark brown. The dorsolateral stripes vanish progressively a short distance posterior to vent.
The head is greyish-brown above with brown vermiculation; upper sides of snout and rostral yellowish-brown,
paler than upper surface; a narrow, dark streak in front of the eye, extending from the nasal to the upper part of the
preocular; supralabials bright ivory; 1
st
–5
th
SL dotted with minute black dots; 2
nd
–5
th
SL narrowly edged with dark
brown, more broadly on hinder margin of 6
th
SL; a conspicuous, broad, blackish-brown postocular streak extends
from behind the eye to the corner of the mouth on the 2 lower postoculars, anterior temporals, upper part of 6
th
SL,
much of 7
th
SL and the whole of 8
th
SL, this latter scale with an elongate bright ivory blotch in its upper middle;
behind the lower half of the 8
th
SL, the blackish-brown postocular streak extends slightly downwards, without
interruption on the neck, and widens on the side of neck, producing the darker colour of the side of the neck,
blotched with the first dorsolateral blotches, before fading out into the greyish-brown colour of the body sides; on
each side, behind the upper half of the 8
th
SL, a bright, conspicuous, oblique ivory streak, reaching the vertebral
row and producing a bright chevron-like marking on the neck, its apex directed backwards and reaching the first
dorsolateral blotch; a short, pale yellow occipital streak; eyes black. The chin and throat are uniformly creamish-
yellow, with a few faint, diffuse greyish-brown spots on the infralabials.
The venter is uniformly creamish-yellow, with outer tips of ventral scales brown and a few medial brown
spots. The subcaudal surface is uniformly creamish-yellow with outer margins of subcaudals narrowly edged with
dark brown; tip of the tail with dark brown spots.
According to Wall (1925), the freshly killed specimen was “blackish-olivaceous dorsally to the edges of the
ventrals. A series of ill-defined light roundish spots on the 5
th
row above the ventrals and the adjacent halves of the
4
th
and 6
th
rows, continued to the base of the tail. Belly yellowish anteriorly, merging to pale salmon posteriorly,
suffused more deeply laterally; with an occasional small dark round spot on the edges of the ventrals. Head
blackish-olivaceous. A short mesial light streak just behind the parietals. A light well-defined V on the nape
beginning behind the gape. First 5 supralabials with black posterior borders. A black postocular stripe to the sides
of the neck. Chin immaculate.”
Variation (based on 10 preserved and 5 living specimens). Habitus. Body elongate, cylindrical; head ovoid,
elongate, distinct from neck; snout average, accounting for 22.7–28.5 % of HL or 1.0–1.3 times as long as eye
diameter, slightly flattened, narrowing anteriorly, blunt when seen from above, rounded seen in profile, with no
defined canthus rostralis; nostril laterally positioned; eye large, its diameter 2.0–2.6 times the distance between its
inferior margin and upper lip edge; pupil round; tail long, cylindrical and progressively tapering.
The maximal total length known is 770 mm (SVL: 520 mm, TaL: 250 mm; male; specimen not preserved from
Arunachal Pradesh). The longest known female is 725 mm long (SVL 531 mm, TaL 194 mm; CAS 215036). Ratio
TaL / TL: 0.262–0.325, with a clear sexual dimorphism (see below).
Dentition. Maxillary teeth: 18–22, with 16–20 subequal teeth + 2 strongly and abruptly enlarged teeth, about
2.0–2.2 times larger than anterior teeth, either without diastema (see Fig. 10) or with a short diastema.
Body scalation. Dorsal scales in 19-19-17 rows, all scales strongly keeled, including those of the 1
st
dorsal
scale row, more strongly keeled on the upper part of the dorsum; all scales strongly and deeply notched apically,
especially on the posterior part of the body; 2 apical pits present. VEN: 158–173 (plus 1 or 2 preventrals); SC:
85–108, all paired; anal plate divided.
Head scalation. Rostral about 1.5–1.6 times as wide as high; nasal rectangular, longer than high, with a lateral
nostril piercing in the middle of the scale; internasals subtriangular, as long as wide or barely longer than wide,
anteriorly narrowed but truncated, with the fore width about 0.45–0.6 times as great as the hind width; prefrontals
about 1.0–1.3 times as long as internasals; one large, longer than wide, undivided supraocular on each side; frontal
bell-shaped, elongate, 1.4–1.7 times as long as wide, 2.2–2.7 times longer than prefrontals; parietals large, in
contact along a suture about 0.9–1.1 times as long as frontal; one subrectangular loreal on each side, 1.1–1.2 times
longer than high; 8 (rarely 9, in 3 / 28 occurrences) supralabials, 1
st
and 2
nd
SL in contact with nasal, 2
nd
and 3
rd
SL
in contact with loreal, 3
rd
–5
th
(rarely 4
th
–6
th
) SL entering orbit, 7
th
SL largest; no subocular; on each side 1 (rarely 2)
preoculars, tall and narrow; 3 small postoculars in all examined specimens; 1 (in 10/28 occurrences) or 2 (in 14/28)
anterior temporals with the formulae 1+2 or 2+2; 9 or 10 (8 and 11 in 1/28 occurrences) infralabials, 1
st
pair in
contact, 1
st
–4
th
or 1
st
–5
th
infralabials in contact with anterior chin shields, 4
th
and 5
th
infralabials largest.
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FIGURE 1. Amphiesma clerki (Wall, 1925), holotype, BMNH 1946.1.13.50. General dorsal view, ventral view and dorsal and
lateral head view. Photographs by Patrick David.
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REVALIDATION OF NATRIX CLERKI WALL, 1925
FIGURE 2. Amphiesma clerki (Wall, 1925), holotype, BMNH 1946.1.13.50 (left) compared to Amphiesma parallelum
(Boulenger, 1890), specimen BMNH 1946.1.13.53 (lectotype). Photographs by Patrick David.
FIGURE 3. Amphiesma clerki (Wall, 1925), live specimen AP1. Photograph by Viral Mistry.
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FIGURE 4. Amphiesma clerki (Wall, 1925), live specimen AP 2. Photograph by Ishan Agarwal.
FIGURE 5. Amphiesma clerki (Wall, 1925), live specimen AP 3. Photograph by Ashok Captain.
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REVALIDATION OF NATRIX CLERKI WALL, 1925
FIGURE 6. Amphiesma clerki (Wall, 1925), live specimen RS09. Photograph by Rohan Pandit.
Colour and pattern. In preservative (Fig. 1–2), the upper body is rather dull, dark greyish-brown, dark
brownish grey, brownish-lead grey, dark grey or dark olivaceous brown, darker than sides of the body at midbody;
body sides are tan, pale brown, chestnut-brown or dark brown but are usually darker on the sides of the neck and
extreme forepart of the body, such as dark chestnut-brown or blackish-brown, distinctly darker than the remaining
part of body sides; this darker lateral colour gradually merges with the lateral colouration but extends over the
entire lateral surface in some very dark specimens; scales of the 1
st
to 4
th
or 5
th
dorsal scale rows broadly edged with
blackish-brown or black, producing dark reticulations and conspicuous irregular, oblique blotches on the sides
arranged in a complex and irregular pattern; dorsum nearly uniform or usually irregularly marked with small, black
spots more or less arranged as series; laterally, a series of distinctly enlarged and conspicuous, irregular, ivory, pale
yellow or pale yellowish-brown blotches on the neck and fore part of body, extending on 4
th
–7
th
or 5
th
–7
th
DSR,
progressively turning into smaller and darker, yellowish-brown or pale brown blotches on the upper half of 5
th
, 6
th
and lower half of 7
th
DSR, rounded and enlarged on the fore quarter to third of the body, more elongate, faint and
reduced to the 5
th
and 6
th
rows posteriorly, connected from the first quarter to third of the body by a more or less
distinct pale or dark yellowish-brown or rusty brown dorsolateral stripe extending along on the 6
th
, more
conspicuous on the hind part of the body; in some specimens, the brown colour on the lateral aspect of the neck and
first quarter of the body is interrupted by pale yellow vertical bars originating from the ventral colour and reaching
up to the levels of the 6
th
or 7
th
DSR, usually much shorter, often reduced to paler areas or absent in most
specimens; scales of the 1
st
dorsal scale row marked with short, irregular blackish-brown streaks, producing a very
discontinuous, zigzag-like ventrolateral stripe, sometimes present only in the hind part of the body. Upper tail
surface as the body, with scales strongly edged with dark brown; dorsolateral stripes vanishes progressively a short
distance posterior to vent.
The head is greyish-brown, brown, brownish-grey above, usually with dark brown vermiculation; upper sides
of snout and rostral paler than upper surface and rather pale brown or dark yellowish-brown; narrow, dark streak in
front of the eye from the nasal to the upper part of the preocular; supralabials ivory, creamish-yellow, yellowish-
brown or pale creamish-brown; 1
st
–3
rd
to 1
st
–5
th
SL dotted with minute brown to black dots or even nearly brown ;
2
nd
–5
th
SL narrowly edged with dark brown, with usually a broader, more conspicuous streak on the hinder margin
of the 6
th
SL; a conspicuous, broad, blackish-brown or black postocular streak extends from behind the eye to the
corner of the mouth, covering lower postocular, anterior and part of posterior temporals, upper part of 6
th
SL, at
least the upper half of 7
th
SL and the whole of 8
th
SL; 8
th
SL uniformly dark or with a pale (ivory or pale yellow)
blotch or irregular in its middle; posterior to the lower half of the 8
th
SL, the blackish-brown postocular streak
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extends without interruption or, in a few occurrences, with a short, heavily dark spotted gap, onto the lateral surface
of neck before fading into the paler background colour of the lateral aspect of the body; posterior to the upper half
of the 8
th
SL, on each side, a bright, conspicuous, oblique ivory, yellow or yellowish-brown streak, reaching the
vertebral row where each branch merges and produces a bright V or Y-shaped chevron-like marking on the neck, its
apex directed backwards and reaching the first dorsolateral blotch; a short, pale yellow occipital streak usually
present. The chin and throat are uniformly ivory, creamish-yellow or pale yellow, with a few faint, diffuse greyish-
brown spots on the infralabials.
The venter is uniformly ivory or creamish-yellow, with outer tips of ventral scales brown or dark greyish-
brown, and a few brown spots on outer parts of some ventrals. The subcaudal surface is as the venter with outer
dark margins of subcaudals narrowly edged with dark brown or black; tip of the tail with dark brown spots or
greyish-brown.
FIGURE 7. Map showing the known distributions of A. parallelum (triangles) and A. clerki (dots).
In life (Fig. 3–6), the upper body is dark olive green to dark greyish-brown with faint black transverse
markings, reticulations and conspicuous irregular, oblique blotches, especially on the forepart of the body; the
dorsal colour pattern is more or less uniform throughout, except in one specimen it is interrupted by reticulation on
the forepart just posterior to the chevron pattern on the nape; the transverse markings are formed by alternate scales
irregularly black edged on the same row; some of the scales along the black transverse marking are bright red; the
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black transverse markings continue on the tail with the marking on the first scale row forming a continuous
transverse line or band which extends onto the tail; interstitial skin white; on each side, a bright red, faint
dorsolateral line extends along the length of the body on both sides on the 6
th
and 7
th
DSR.
Dorsal sides of the head are dark olive green to greyish brown; a faint pale brown streak on the centre of the
nape and occiput; a black or blackish-brown preocular and postocular streak extends from the tip of the anterior
nasal scale through the eye and, posteriorly, towards the extremity of the nape where it fades into the paler body
colour; supraoculars with a cream or yellow streak that extends on the top of the nape as a distinct line and merges
with each other forming a Y- or V-shaped chevron marking on the nape; supralabials bright yellow or sometimes
white, speckled with black. Chin and throat are bright yellow or sometimes white. The iris is dark brown or black
with a bright red ring around it; other parts of the eye dark olive green to dark grey.
The venter is bright yellow to white; outer margins of ventral scales bright red with black markings; the bright
red blotches continuing to the tip of the tail.
Sexual dimorphism. It is marked in two characters: (1) ratio tail / total length: males: 0.280–0.325 vs. females:
0.262–0.273; and (2) the number of subcaudals: males: 87–108 vs. females: 84–85. Quite interestingly, the small
diastema of maxillary teeth was seen only in females.
Distribution (Fig. 7; numbers refer to the circles on the map). This species is known only from the eastern
part of the Himalaya Range and adjacent ranges. Nepal (?). BMNH 58.6.24.5, “Nepal” (1). India. Sikkim
. No
precise locality (2). West Bengal
. Darjeeling (3). Arunachal Pradesh. West Kameng District. Eaglenest Wildlife
Sanctuary (4); Lower Subansiri District. Talle Valley, near Hapoli (5); Changlang District. Outskirts of Shidi
(Gandhigram) (6). Nagaland
. Kohima District. Sechu (7). Myanmar. Kachin State. Vicinity of Watamkawng (8);
Sinlumkaba (9). People’s Republic of China. Yunnan Province
. Nu Jiang Nature Reserve, near Pianma, Nu Jiang
County (10).
The locality of specimen BMNH 58.6.4.25 is given as “Nepal”, without precision. It was deposited by B. H.
Hodgson, who collected in Nepal but also in Sikkim. Additional specimens from Nepal are necessary to confirm
the occurrence of A. clerki in that country; see also the discussion given below under the account of Amphiesma
parallelum and the discussion given in Vogel & Hauser (2006) on the Nepalese locality of Ptyas nigromarginata
(Blyth, 1854).
Comments. This species shows consistent morphological differences with Amphiesma parallelum (Boulenger,
1890) that are detailed below in the Discussion.
Amphiesma parallelum (Boulenger, 1890)
(Fig. 8–9)
Tropidonotus parallelus Boulenger, 1890: 345. Type locality. By virtue of lectotype designation: “Sikkim”, currently the State
of Sikkim, India. Lectotype. By designation of Kramer (1977: 728): BMNH 1946.1.13.53, adult male; collected and
deposited by Sir Joseph Dalton Hooker.
Material (n = 18 + 1 unpreserved specimen). India. Sikkim (? See below). BMNH 1946.1.13.53 (lectotype of
Tropidonotus parallelus Boulenger, 1890), “Sikkim”.—Meghalaya
. BMNH 1946.1.12.83–84,
BMNH 1946.1.13.48 (3 former syntypes of Tropidonotus parallelus Boulenger, 1890), “Khasi Hills”; ZSI 3852,
“Shillong”; ZSI/ERS 112, ZSI/ERS 205, ZSI/ERS 2785, ZSI/ERS 3076–3077, Risa Colony, Shillong; ZSI/ERS
272, ZSI/ERS 9059, Tripura Castle Road, Shillong; ZSI/ERS 450, ZSI/ERS 970, Fruit Garden, Shillong; ZSI/ERS
3253, Mawlai, East Khasi Hills District; ZSI/ERS 8262, Mawphlang, East Khasi Hills; ZSI/ERS 9060, Selbelgiri,
Garo Hills.—Nagaland
. Unpreserved specimen (Fig. 9), near the Tragopan Sanctuary (25.63549N-094.01261E),
Khonoma.—No locality
. ZSI 4397, “Madras Hills”, obviously in error.
Systematics. Boulenger (1890) did not specify the number of syntypes upon which he based his description
but he mentioned the type locality as “Sikhim [sic], Khási Hills, hills of Upper Burma and Yunnan”. Subsequently,
Boulenger (1893: 223) considered six specimens to be syntypes, i.e. one from Sikkim (BMNH 1946.1.13.53), three
from the Khasi Hills (BMNH 1946.1.13.48 and BMNH 1946.1.12.83–84), one from Sanda, in northern Myanmar
(BMNH 1946.1.21.87) and the last one from Hotha Valley, Yunnan (BMNH 1946.1.13.58). These two latter
specimens were collected by J. Anderson. We examined them and they are referable to Amphiesma bitaeniatum
(Wall, 1925). The four other original type specimens, all are referable to the same species that should now be
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known as A. parallelum. Kramer (1977: 728) considered specimen BMNH 60.3.19.1359, from Sikkim but not
collected by Sir J. Hooker, to be also a syntype of Tropidonotus parallelus. This assertion is erroneous, as this
specimen did not belong to the original type material. However, quite interestingly, Kramer noted that this
specimen disagreed in several points with the other syntypes, which possibly represented two forms. This author
was correct, as specimen BMNH 60.3.19.1359 is referable to Amphiesma clerki (see above). In order to fix the
status of the taxon Tropidonotus parallelus Boulenger, 1890, Kramer (1977) designated the specimen BMNH
1946.1.13.53, also from Sikkim, as its lectotype.
FIGURE 8. Amphiesma parallelum (Boulenger, 1890), specimen BMNH 1946.1.12.83 (top) and specimen BMNH
1946.1.12.84 (bottom). Views of left sides of heads. Photographs by Patrick David.
However, in our opinion, there remains an ambiguity on the exact collection locality of the lectotype. To our
best knowledge, no other specimen of A. parallelum has ever been collected in Sikkim (Sanyal et al. 2006) or in the
adjacent State of West Bengal. Specimen BMNH 1946.1.13.53 was collected and deposited by Sir Joseph Dalton
Hooker (1817–1911), one of the greatest botanists and explorers in Asia of his time. In 1847, he undertook an
expedition to the Himalaya Range which lasted four years. Hooker travelled extensively in the region of Darjeeling
and, during nearly the whole of 1849, in Sikkim where he gathered a large collection, mostly of plants. However,
Sikkim was not his sole area of interest in India. He spent much of 1850 in Assam, especially in the Khasi Hills
(Hooker 1854; Turrill 1963; Desmond 2006). Unfortunately, his journal does not contain much information on the
snakes collected during his trips. In Vol. 2 (p. 25–26), Hooker stated to have collected “about 12 species” in
Sikkim, of which seven were colubrid snakes, without additional details. Moreover, in the same volume (Hooker
1854: 305) wrote “Reptiles, and especially Colubridae, are very common in the Khasia mountains, and I procured
sixteen species and many specimens. The natives repeatedly assured us that these were all harmless, and Dr. Gray,
who has kindly examined all my snakes, informs me of the remarkable fact (…) that whereas none are poisonous,
(…).” Obviously, Hooker also collected numerous colubrid snakes in the Khasi Hills. Although proof is lacking,
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we cannot exclude a confusion in specimen labels and localities. Instead of Sikkim, the lectotype specimen might
well come from the Khasi Hills, where numerous specimens of Amphiesma parallelum have been subsequently
collected.
Amphiesma parallelum has been extensively confused in the literature with A. clerki but also with A.
bitaeniatum and A. octolineatum, as well as with A. platyceps and A. sieboldii. A chresonymy of A. parallelum,
however, is out of the scope of the present paper. Such confusion appeared in major authors such as Bourret (1936)
and Smith (1943). As currently conceived, Amphiesma parallelum is a monotypic species.
As far as we know, Amphiesma parallelum had never been depicted alive in the literature before the present
paper. Only a colour painting appeared in Das (2010: Pl. 67: Fig. 13).
FIGURE 9. Amphiesma parallelum (Boulenger, 1890), live, dorsal and ventral view, specimen from Nagaland. Photograph by
Viral Mistry.
Diagnosis. A species of the genus Amphiesma characterized by (1) a dark dorsal background in life, dark
reddish-brown or probably dark chestnut-brown in life, turning to cream, grey, pinkish-grey, tan, more or less dark
greyish-brown or dark yellowish-grey in preservative, (2) on each side, a conspicuous brown or reddish-brown
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(cream, pale greyish-brown or pale yellowish-brown in preservative) dorsolateral stripe, narrowly edged on its both
sides with dark brown or black, extending on the 5
th
–7
th
dorsal scale rows from the occipital region along the whole
of the body, (3) many dorsal scales longitudinally edged with dark brown, especially on the lower part of the sides,
producing a faint reticulation or irregular streaks, (4) upper edge of scales of the 1
st
dorsal scale row and lower edge
of those of 2
nd
row blackish-brown, producing a very irregular ventrolateral stripe, (5) a rather narrow, irregular,
blackish-brown postocular streak extending from behind the eye to the corner of the mouth, usually separated by a
wide gap from the dark line edging the lower side of the dorsolateral stripe, (6) no chevron on the upper part of the
neck, (7) supralabials mostly uniform, not (or barely) spotted, (8) venter uniformly cream, beige or yellowish-
brown, with the outer edge of ventral tips blackish-brown, (9) 21–22 maxillary teeth, the last two moderately
enlarged and separated from anterior teeth by a distinct diastema, (10) 19–19–17 dorsal scale rows, (11) dorsal
scales moderately to strongly keeled on 2
nd
–9
th
dorsal and vertebral scale rows, smooth or at best feebly keeled 1
st
dorsal row, (12) tail moderate, proportion tail length / total length 0.221–0.252 (0.239–0.252 in males, 0.221–0.249
in females), (13) 160–173 ventrals, (14) 63–77 subcaudals, (15) anal plate divided, (16) usually 2 preoculars, and
(17) 1 or 2 anterior temporals.
Variation (based on 18 preserved and 1 living specimens). Habitus. Body elongate but rather plump in
females, cylindrical; head ovoid, elongate, distinct from the neck; snout length average, accounting for
19.7–23.1 % of HL or 1.1–1.2 times as long as eye diameter, flattened, narrowing anteriorly, blunt when seen from
above and in profile, with no defined canthus rostralis; nostril laterally positioned; eye size average, its diameter
1.6–2.0 times the distance between its inferior margin and upper lip edge; pupil round; tail length average,
cylindrical, and progressively tapering.
The maximal total length known is 633 mm (SVL: 483 mm, TaL: 150 mm; female; specimen ZSI/ERS 9070).
The longest known male is 601 mm long (SVL 450 mm, TaL 151 mm; ZSI/ERS 272). Proportion TaL / TL:
0.221–0.252, without clear sexual dimorphism.
Dentition. Maxillary teeth: 21–22, with 19–20 subequal teeth + 2 moderately enlarged posterior teeth, not
twice as long as an anterior teeth, separated from the anterior teeth by a distinct diastema, at least as wide as the
length of a posterior tooth.
Body scalation. Dorsal scales in 19-19-17 rows, scales of the 2
nd
–9
th
rows strongly keeled and deeply notched
at their apical part, especially on the posterior part of the body, more strongly keeled on the upper part of the
dorsum; scales of the 1
st
dorsal row enlarged, smooth or feebly keeled; no apical pit. VEN: 160–173 (plus 1 or 2
preventrals); SC: 63–77, usually all paired but some single scales has been recorded in a few specimens; anal plate
divided.
Head scalation. Rostral about 1.5–1.6 times as wide as high; nasal rectangular, longer than high, nostril lateral
and piercing in the middle of the scale; internasals subtriangular, 0.9–1.1 times as long as wide, anteriorly narrowed
but truncated, with the fore width about 0.45–0.55 times as great as the hind width; prefrontals about 1.0–1.3 times
as long as internasals; 1 large, longer than wide undivided supraocular on each side; frontal bell-shaped, rather
small, 1.3–1.5 times as long as wide, 2.2–2.7 times longer than prefrontals; parietals large, in contact along a suture
about 0.8–1.2 times as long as frontal; one subrectangular loreal on each side, as long as high or barely longer than
high; 8 supralabials in all examined specimens, 1
st
and 2
nd
SL in contact with nasal, 2
nd
or 2
nd
and 3
rd
SL in contact
with loreal, 3
rd
–5
th
SL entering orbit, 7
th
SL largest; no subocular; on each side 2 (exceptionally 1) preoculars, small
and narrow; 3 (rarely 2) small postoculars; 1 (in 17/34 occurrences) or 2 (in 17/34) anterior temporals with the
formulas 1+1, 1+2, 2+2 or rarely 2+3 temporals; 9 or 10 infralabials, 1
st
pair in contact, 1
st
–4
th
or 1
st
–5
th
infralabials
in contact with anterior chin shields, 4
th
and 5
th
infralabials largest.
Coloration and pattern. In preservative (Fig. 8), the body is cream, pinkish-grey, tan, pale to dark greyish-
brown, dark yellowish-grey or even dark brown, not or barely darker on the upper part of the body than on the
sides; many scales of the sides and upper dorsum surface either narrowly and irregularly edged with blackish-
brown, especially on the sides, or paler centred, i.e., pale yellowish-grey or yellowish-brown, or both, producing an
irregular, diffuse reticulate pattern or dark elongate streaks; scales more broadly edged or mottled with dark brown
or blackish-brown a short distance just behind the head, where sides can be irregularly but distinctly darker than
other part of the sides; scales of the 1
st
dorsal scale row more largely pale centred; upper dorsal surface sometimes
with scattered black spots; on each side, a broad, conspicuous dorsolateral stripe, cream or pale greyish-brown,
anteriorly, slightly larger, i.e. yellowish-grey or pale yellowish-brown after the anterior quarter of the body, and
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narrowly edged with a dark brown or black line on both its lower and upper sides, extends on the upper part of 5
th
,
the whole of 6
th
and much of 7
th
dorsal scale rows from the nuchal region behind temporals scales, along the whole
of the body; this dorsolateral stripe is narrowly but distinctly edged with irregular, discontinuous lines, more
conspicuous on the forepart of the body, the lower one composed of dark brown or blackish-brown streaks on the
upper half of 5
th
DSR, the upper one by similar streaks on the upper part of scales of the 7
th
DSR; upper halves of
scales of the 1
st
DSR and often lower part of those of the 2
nd
DSR row marked with short, irregular blackish-brown
streaks, producing a very discontinuous stripe, sometimes present only in the hind part of the body or barely
noticeable; lower part of scales of 1
st
with irregular blackish-brown streaks, producing a very discontinuous,
zigzag-like ventrolateral stripe. Upper tail surface as the body but, anteriorly, somewhat paler on its upper
dorsolateral half due to the continuation of the pale dorsolateral stripe at least to the fore part of the tail, stripes
separated by dorsal background colour on a narrow vertebral area; scales of the lower part of the sides rather pale,
distinctly edged with dark brown, strongly edged for the scales of the 1
st
row, producing a strongly reticulate
pattern; hind half of tail uniformly pale as the dorsolateral stripe with scales strongly edged with brown.
The head is beige, pale greyish-brown, yellowish-brown or yellowish-grey above, distinctly paler than body in
dark specimens; sides of the snout, supralabials and temporal areas ivory, creamish-yellow or pale creamish-
brown; upper head scales irregularly and incompletely edged with dark brown; a short, narrow, dark streak in front
of the eye on the upper edge of lower preocular and loreal, not reaching nasal and often reduced to an elongate
blotch between the two preoculars; supralabials entirely uniform or with, at most, a few dark dots, not edged with
dark brown; a narrow, oblique blackish-brown or black postocular streak extends from behind the eye to the corner
of the mouth, covering lower postocular, lower half of anterior temporal, upper edge of 6
th
SL, the upper half of 7
th
SL and lower half of 8
th
SL; the postocular streak does not extend beyond 8
th
SL or at most only a short distance,
and is usually widely separated from both the dark line edging the lower side of the dorsolateral stripe and, if
present, the darker colour of the body side, by a pale coloured gap of same colour as the supralabials and chin;
sometimes this gap is much reduced, with postocular streak and dark body colour nearly in contact; the pale nuchal
gap above and behind the end of the postocular streak is edged with blackish-brown, more broadly on its lower
edge, and merges into the dorsolateral stripe, the dark edges of the gap extending as edges of the dorsolateral stripe;
a cream or pale yellow occipital streak, narrowly black-edged, extends from the tip of parietals to the upper part of
the nuchal pale-coloured gap. The chin and throat are uniformly ivory, creamish-yellow or pale yellow; some
infralabials with faint, darker edges.
The venter is uniformly ivory, creamish-yellow or pale yellowish-brown, with outer edges of tips of ventral
scales dark brown or blackish-brown, in contact with the short streaks of the lower edges of scales of the 1
st
DSR.
The subcaudal surface is uniformly coloured as the venter.
In life (Fig. 9), the colouration is much more vivid than in preserved specimens. According to the sole
available specimen, the dorsum is dark reddish-brown with many dorsal scales irregularly black-edged at random
locations, not forming any definite pattern; on each side, a pale brown dorsolateral stripe extending on the 6
th
and
7
th
dorsal scale rows from the nape to the end of the body; these two stripes appear to be truly parallel; on each side
of the pale dorsolateral stripe, dorsal scales are regularly black-edged, producing a thin, regular black lines above
and below the stripe; these black lines extend to the end of the tail dorsolaterally; tail as the body, with some of the
lateral scales with a red centre.
The head is dark brown above, brighter reddish-brown on the nape; an irregular, thin, black preocular streak
starts form the anterior nasal scale to the eye; supralabials, lower preocular and loreal white marked with irregular
red and black spots; the last supralabial is reddish-brown; a thin postocular streak from the eye to the gap behind
the corner of the mouth, extending on penultimate supralabial; last supralabial brown; chin and throat white with
black and red markings.
The venter is pale yellow; outer tips of ventral scales with red spots, plus other black spots at the intersection of
the pale yellow background colour and red spots, producing an irregular ventral red stripe dotted with black, that
extends venterolateraly from the throat to the vent. Subcaudal surface pale yellow, progressively merging with the
lateral red spots that extends to the tip of the tail.
Sexual dimorphism. It is weakly expressed only in the number of subcaudals: males: 72–77 (n = 8; x = 74.3; s
= 1.6) vs. females: 63–73 (n = 8; x = 70.0; s = 3.8).
Distribution (Fig. 7; numbers refer to the triangles on the map). As currently conceived, Amphiesma
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parallelum is definitely known only from mountain ranges (Khasi Hills, Garo Hills and Mts. Naga) south of the
Himalaya Range. India. Sikkim
. Presence unconfirmed, perhaps in error (1). Meghalaya. Shillong and its vicinity
(2); Mawphlang, East Khasi Hills (3); Mawlai, East Khasi Hills District (4); Selbalgiri, Garo Hills (5). Nagaland
.
Tragopan Sanctuary (25.63549N-094.01261E), Khonoma (6).
Lastly, we examined specimen BMNH 58.6.24.5, alledged to come from “Nepal”. According to its pattern, it is
indeed an Amphiesma clerki (see above).
FIGURE 10. Maxilla of specimen RS09, Amphiesma clerki (Wall, 1925). Note the absence of diastema between anterior teeth
and the strongly and abruptly enlarged posterior teeth. Photograph by Viral Mistry.
Discussion
A summary of differences between Amphiesma parallelum and A. clerki
In spite of their long confusion, Amphiesma parallelum and Amphiesma clerki can be rather easily separated by
a series of contrasting morphological characters including aspects of pattern, morphometry, scalation and dentition.
Differences in these last three series of characters are summarized in Table 1. A comparative summary of
colouration and pattern between the two species can be established as follows:
Amphiesma clerki: (1) a dark dorsal background in life and preservative with an indistinct dorsal pattern; (2) a
series of pale dorsolateral blotches and spots on each side, much larger and conspicuous on the neck and forepart of
the body; (3) a pale and faint dorsolateral stripe, often absent on the forepart of the body; (4) irregular dark blotches
on the sides; (5) pale blotches on the sides of the neck; (6) a broad postocular streak extending from behind the eye
to the side of the neck usually without gap (rarely discontinuous); (7) a conspicuous, V or Y-shaped pale chevron
on the neck; (8) supralabials dotted with dark dots or even nearly brown, and edged with dark brown; (9) outer tips
of ventral scales dark.
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Amphiesma parallelum: (1) a dark dorsal background in life, much paler in preservative with a well-defined
pattern; (2) no pale dorsolateral spots or blotches; (3) a conspicuous dorsolateral stripe, narrowly edged with dark
brown or black, extending on the from the occipital region along the whole of the body; (4) no large dark blotches
on the sides replaced by faint reticulations or irregular streaks; (5) no pale blotches on the sides of the neck; (6) a
narrow postocular streak extending from behind the eye to the 8
th
SL, separated from the dark colour of the side by
a distinct pale gap, sometimes reduced to a small area; (7) no chevron on the neck; (8) supralabials mostly pale and
uniform; (9) outer tips of ventral scales not dark, or only on their edge.
Comparison with other species of the Amphiesma parallelum group
A revision of the group of Amphiesma parallelum is out of the scope of the present paper but we felt it
necessary to address the differences between species of the group. Besides A. clerki, A. parallelum has also been
confused with Amphiesma bitaeniatum (Wall, 1925) and A. octolineatum (Boulenger, 1904). Morphometric and
meristic characters are summarized in Table 1.
TABLE 1. Comparison between main morphological characters separating A. clerki and A. parallelum, and other species
of the A. parallelum group.
continued.
Characters of A. platyceps and A. sieboldii are drawn from our material and Malnate (1966); those of A. metusia are
from Zhao et al. (1998) and Zhao (2006).
However, in the the genus Amphiesma, it is more convenient to identify these species by their dorsal and
cephalic patterns. These four species share a grey, greyish-brown, yellowish-brown or pale brown background
dorsal colour and striped pattern with at least one paler, more or less distinct, black edged dorsolateral stripe on
each side, to which we add a combination of other characters defined below. Our data show that there is no overlap,
or barely significant, between the combination of pattern characters among these species.
We retained the following characters in pattern and coloration that proved to be diagnostic either alone or in a
combination: (A) general dorsal pattern, (B) number of dorsal stripes, (C) difference of tone between the dorsum
and sides of the body, (D) width of the lower black edge of the main dorsolateral stripe, (E) continuity of the
postocular streak with one of the black lateral dorsolateral stripes, (F) width of the ventrolateral stripe, and (G)
ventral pattern. Results are given in Table 2.
The differences in the darkness of the middorsal and lateral surfaces can be easily used to distinguish A.
parallelum from A. clerki, and these species from A. bitaeniatum, A. octolineatum and A. metusia. The significance
Species Tooth diastema TaL/TL Ven SC
A. clerki absent or short 0.265–0.325 158–173 85–108
A. parallelum wide 0.221–0.252 160–173 63–77
A. bitaeniatum absent 0.242–0.273 153–177 76–88
A. octolineatum absent 0.209–0.264 152–177 57–91
A. metusia absent 0.234–0.280 159–164 72–85
A. sieboldii wide 0.242–0.301 168–207 81–111
A. platyceps wide 0.232–0.286 191–234 78–98
Species KSR K1SR NSR Preoc ATe
A. clerki +++ +++ +++ 1 (2) 1–3
A. parallelum ++/+++ 0/+ 0/+ 2 (1) 1–2
A. bitaeniatum ++/+++ 0 +++ 1 (2) 1–2
A. octolineatum +001 (2)1–2
A. metusia ++ 0011–2
A. sieboldii ++ 0/+ 0/+ 1 (2) 1–2
A. platyceps 0/+ 0011–2
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in the continuity or discontinuity of the postocular streak with the lateral black stripe of the body (Character E),
already noticed by Wall (1925), is confirmed.
TABLE 2. A comparison of main distinctive characters of dorsal patterns in members of the Amphiesma parallelum-
group.
Characters are as follows: (A) general dorsal pattern: (1) only striped, or (2) striped and blotched, or (3) only blotched.
(B) number of dorsal stripes: (0) none, (1) only one, or (2) at least two on each side (not including the ventrolateral
stripe). (C) difference of tone between the dorsum and sides of the body: (1) equal or subequal, (2) sides distinctly darker
than the dorsum, (3) dorsum distinctly darker than the sides. (D) width of the lower black edge of the main dorsolateral
stripe: (0) no dorsal stripe, (1) narrow, line-like, or (2) broad, stripe-like. (E) continuity of the postocular streak with one
of the black lateral dorsolateral stripes: (0) no dorsal stripe, (1) no gap, i.e. continuous from the corner to the mouth to the
side of the body, or (2) a distinct, wide gap at the level of the corner of the mouth and fore part of the neck, sometimes
reduced to a narrow gap making both parts of the black stripe nearly in contact. (F) width of the ventrolateral stripe: (1)
narrow, line-like, or (2) broad, covering a significant width of scales of the 1
st
dorsal scale row. (G) ventral pattern: (1)
immaculate, with tips of ventrals dark-edged, or (2) with distinct spots or blotches on outer parts of ventrals, or (3)
throughout dotted or speckled.
A comparison bearing on scalation and morphometry between species of the A. parallelum-group is given in
Table 1. Variation in Amphiesma parallelum is based on our own data, and not on literature. In considering
morphometric, dentitional and scalation characters, species of the Amphiesma parallelum group can be separated
by the following key. Morphological characters are drawn from our material and from references cited above for
Table 1.
1A No dorsolateral stripe, sometimes a row of aligned white dots; a narrow, black or dark brown preocular and postocular streak;
more than 190 ventrals in males . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1B At least one broad, pale dorsolateral stripe extending from neck to the end of tail or at least present on the hinder part of the
body; a conspicuous, black or dark brown postocular streak, preocular streak present or absent; less than 178 ventrals . . . . .3
2A Only upper dorsal scale rows moderately keeled, others smooth; 205–234 ventrals in males, 191–216 VEN in females; dorsal
surfaces usually marked with scattered black dots, without a dorsolateral series of white dots (or barely defined); venter usually
immaculate; subcaudal surfaces usually immaculate, rarely darkened with grey. . . . . . . . . . . . . . . . . . . . Amphiesma platyceps
2B All dorsal scale rows (excepted 1
st
one) distinctly keeled; 191–207 VEN in males, 168–190 VEN in females; dorsal surfaces
usually uniformly brown, except a dorsolateral series of well-defined small white spots; venter often largely speckled with
dark greyish-brown posteriorly; subcaudal surface usually dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma sieboldii
3A Dorsal pattern only striped, either with one single dorsolateral stripe or several stripes on each side, without large blotches or
spots on the sides (black spots possibly present on the back) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3B Dorsal pattern both striped and blotched, with the combination of a single dorsolateral stripe, dark dark spots and blotches on
the sides, and dark spots on the back, producing a rather irregular, complex general pattern . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
4A A single, broad, conspicuous pale dorsolateral stripe on each side, not narrowing on the nape; dorsum darker than body sides or
similar in tone; dorsal scales strongly keeled and more or less notched. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4B Several stripes on each side: main dorsolateral stripe bordered below with one or two dark grey or dark brownish-grey stripes,
the lower one, irregular, very wide, making the lateral aspect distinctly darker than the dorsum; pale dorsolateral stripe ending
anteriorly in narrowing on the nape; dorsal scales smooth or weakly keeled, not or very slightly notched. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma octolineatum
5A Body cream, grey or in various shades of brown in preservative (dark reddish-brown in life), with sides and upper dorsal sur-
face of same tone; a conspicuous pale dorsolateral stripe, edged above and below with a narrow dark line; a narrow, black
postocular streak separated from the dark line edging the lower side of dorsolateral stripe by a distinct gap on the neck side; a
dark preocular streak present; enlarged posterior maxillary teeth separated from others by a distinct diastema; 63–77 subcau-
Characters
Species
ABCDE F G
A. clerki 21311 (rarely 2)12
A. parallelum 11112 11
A. bitaeniatum 11321 22
A. octolineatum 12221 22
A. metusia 22101 01
A. sieboldii 30100 03
A. platyceps 30100 01
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REVALIDATION OF NATRIX CLERKI WALL, 1925
dals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma parallelum
5B Body dark grey, dark brown or dark brownish-yellow above, pale brown or yellowish-brown on the sides, distinctly darker
dorsally than laterally; a conspicuous pale dorsolateral stripe, edged above by a narrow, discontinuous dark line, and below by
a broad dark, thick stripe, sometimes covering up to the upper half of body sides, broader on the neck sides; a broad, black pos-
tocular streak extends on the neck and merges with the lateral dark stripe without gap or constriction; no preocular streak pres-
ent; no diastema between enlarged posterior maxillary teeth and other teeth; 76–88 subcaudals . . . . . Amphiesma bitaeniatum
6A Dorsal colour dark brown, dark greyish-brown or dark brownish-grey; no dorsolateral stripe visible on the fore part of the
body; a series of large, pale spots (reddish-brown or rusty red in life) on the neck and forepart of the body, connected along the
hinder half of the body by a pale dorsolateral stripe; a conspicuous, pale, V or Y-shaped nuchal chevron; scales of the 1
st
DSR
strongly keeled; 85–108 subcaudals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma clerki
6B Dorsal colour dark grey or blackish-brown, heavily checkered or reticulate with black; a poorly defined, pale dorsolateral
stripe, usually interrupted, ending anteriorly in widening on the nape; no pale blotches anteriorly; no nuchal chevron; scales of
the 1
st
DSR smooth; 72–85 subcaudals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Amphiesma metusia
Conclusion
No attempt has been done to establish precise relationships between species referred to the A. parallelum group.
However, it appears that Amphiesma clerki shares characters of both Amphiesma parallelum and Amphiesma
sieboldii (Günther, 1860), as defined for this latter species by Malnate (1966). For example, by its dentition,
especially the greatly and abruptly enlarged posterior maxillary teeth, nearly twice as long as other maxillary teeth,
and the series of dorsolateral spots instead of a stripe, Amphiesma clerki shows clear affinities with Amphiesma
sieboldii. Posterior maxillary teeth are also enlarged in A. parallelum, but to a lesser extent than in the two other
species. In these three taxa, the number of teeth (19–23 + 2–3 enlarged teeth) is similar. However, we observed the
presence of a distinct diastema in A. sieboldii and A. parallelum, whereas the posterior teeth are not, or at best
barely separated from the anterior ones in A. clerki. In contrast, the typical pattern of the neck and the strongly
keeled and notched dorsal scales of A. clerki are notably different from characters found in Amphiesma sieboldii
(Malnate 1966). From A. parallelum, A. clerki is mostly distinguished by the pattern of the body and especially of
the neck, the dentition, the keeling of dorsal scales, and the numbers of ventrals. Lastly, as A. sieboldii, A. clerki is
definitely a high montane species. Specimens from Bompu (Arunachal Pradesh, India) were collected between
1,780 and 2,400 m asl. Although long confused, A. clerki and A. parallelum may not be closely related taxa.
The revalidation of Amphiesma clerki adds one more species to the snake faunas of India, Myanmar and China.
This species is now definitely known only from mountainous areas of northeast India (Sikkim, West Bengal,
Arunachal Pradesh, and Nagaland), northern Myanmar and extreme south-western China (western Yunnan). Its
occurrence in Nepal is possible, if not likely. In contrast to long enduring, erroneous data cited in the literature,
Amphiesma parallelum inhabits also Northeast India, but it is definitely known only from the ranges of the States
of Meghalaya and Nagaland. As explained above, its occurrence in Sikkim is unconfirmed and is perhaps in error.
Amphiesma parallelum has often been cited from Myanmar and China, but all specimens examined by us, except
the specimen of A. clerki cited above, proved to be either A. sieboldii or A. bitaeniatum (see below in the
Appendix). Lastly, David et al. (2005) showed that records of Natrix parallela or A. parallelum from Vietnam, for
example by Boulenger (1890), Bourret (1936) and Nguyên & Hô (1996), refer to Amphiesma bitaeniatum.
More specimens from montane areas of the Eastern Himalayas are vital to ascertain variation and distribution of
Amphiesma parallelum, Amphiesma clerki and other members of this morphologically diverse natricid group.
Acknowledgements
We thank the following people for their help: Ashok Captain (Pune) for data and photographs of the living
specimens from Gandhigram, Arunachal Pradesh, Rohan Pandit (Pune) for his help with collecting the specimen
RS09 and its photographs when in life. VM would like to thank the Nagaland Biodiversity & Conservation
program (2010) for providing the opportunity to visit Nagaland for biodiversity survey during which he found live
Amphiesma parallelum.
It is also a pleasure to thank the following persons who made this review possible in sending us specimens,
providing us data, or letting us examine preserved specimens under their care (in alphabetical order of their
museum and institution acronym): David A. Kizirian and Linda S. Ford (AMNH) Colin J. McCarthy and Patrick
DAVID ET AL.
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Campbell (BMNH), Jens V. Vindum and Alan E. Leviton (CAS), Yuezhao Wang, Xiaomao Zeng, Jiatang Li and
Ermi Zhao (CIB), the late Jarujin Nabhitabhata (CTNRC), Harold Voris and Alan Resetar, (FMNH), Andreas
Schmitz (MHNG), Alain Dubois, Ivan Ineich et Annemarie Ohler (MNHN), Heinz Grillitsch, Franz Tiedemann,
Silke Schweiger and Richard Gemel (NMW), Bob Murphy (ROM), Jianqiang Cen (SMNH), George Zug, Kenneth
Tighe and Ronald Heyer (USNM), Mark-Oliver Roedel and Frank Tillack (ZMB), and Channakesava Murphy
(ZSI, Kolkata).
We thank the State Forest Department, Arunachal Pradesh, for permission to work in Eaglenest. The work in
Arunachal was funded by grants from Rufford Small Grants and Whitley Fund for Nature.
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APPENDIX. Additional examined specimens.
Amphiesma bitaeniatum (n = 21). Myanmar. AMNH 48468, “Huton, Kachin Hills, Burma”, now Huton, Kachin State;
BMNH 1925.9.17.3, BMNH 1925.12.22.19, “Huton, Kachin Hills, Burma”, now in Kachin State; BMNH 1946.1.13.58,
“Kutkai, North Shan States, Burma: 6000 feet”, now Kutkai, Shan State. – People’s Republic of China. Yunnan Province
.
BMNH 1946.1.13.56, “Hotha Valley, Yunnan”, now the valley around Husa, Longchuan County; ZSI 8532, ZSI 8534–36,
ZSI 8540, “Hotha, Yunnan”, now Husa, Longchuan County; BMNH 1946.1.21.87, “Sanda, Upper Irrawady”, now
Lianghe County; CAS 215037, Nu Jiang Nature Reserve near Pianma, on western slope of Gaoligongshan (26°00'03.2N-
98°39'41.6E), ca 7,800 ft, Nu Jiang Xian; SMNH 1259, Yunnan; ZMB 28951, no precise locality; ZSI 8531, “Muangla
Valley”, now Muangla, Lianghe County; ZSI 8539, “Ponsee”, now Pengxi, Yinjiang County. Guangxi Autonomous
Province. 1 specimen, M.W. L
AU
’
S
collection (no number), Cenwanglao Shan. – Thailand. THNHM 25698 (ex
CTNRC 980506), Doi Inthanon, Chiang Mai Province.—Vietnam. Lào Cai Province
. BMNH 1930.11.16.5, “Fan-Si-Pan,
Lao Kay, Tongking”, now Mt. Phang Si Pang; MNHN 1999.9090, vicinity of Sapa; ROM 38098, Lao Cai.
Amphiesma metusia (n = 10). People’s Republic of China. Sichuan Province
. BMNH 1911.12.19.1, “Szechuan”; CAS
195196–195197, vicinity of elev. 2400 m, 9.5 km north of Tuowu (28°49'N - 102°17'E), on the Hanyuan to Xichang Road,
then 1.4 km NNE of dirt road, Liangshan Yizu Autonomous Prefecture; FMNH 18722, “Hsiao Yang Chi”, Sichuan;
FMNH 170647, “Sikang”, now eastern Sichuan; FMNH 232805, 9 km west of Bin Ling, Wa Shan Camp, Hongya Xian;
FMNH 232806, Hongya Xian; USNM 69926–69927, near Washan; ZMB 27866, Washan.
Amphiesma octolineatum (n = 52). People’s Republic of China. Yunnan Province
. AMNH 21022, AMNH 21024, “Lichiang-
fu, 8500 ft”, now Lijiang Naxizu Zizhixian; AMNH 21050, AMNH 21051, “Yunnan: Tengyueh”, now Tengchong County;
AMNH 35210, “Yunnan: Hsin Kai”, Yunnan Province; AMNH 66653, “Yunnan: Kunming”; BMNH 1904.11.29.16–20,
“Ku-taing Fu”, now Gudong; BMNH 1905.1.30.62, “Tongchuan-fu, Yunnan”, now Dongchuan County;
BMNH 1905.5.30.16–20, BMNH 1946.1.12.60, BMNH 1946.1.13.46, BMNH 1946.1.13.57, “Yunnan Fou”, now
Kunming; CAS 64272, “Yunnan”; NMW 22486:1–13, “Yungning, Yung Pe, Likiang, Yunnan, S. China”now in Lijiang
Naxizu Zizhixian; ZMB 65438–654441, ZMB 65571, ZMB 65576, ZMB 65579, ZMB 65582–655584, “Talifu W-
Yunnan”, now Dali County. Sichuan Province
. MNHN 1912.0267–0271, “Mienning (2000 mètres)”, now Mianning
County (28°35’N-102°11’E). Guizhou Province
. SMNH 2527, Yin River, Fanjing Shan, Hengyuanzi, 1,800 m. – No
precise locality. MNHN 1905.0289, MNHN 1907.0012, “Chine”.
Amphiesma platyceps (n = 8). India. Jammu and Kashmir
. MNHN 1988.6484, above Doda, between Makambagi and
Ularbagi, Udamphur District, at about 2,800 m; ZMB 7293, “Kashmir”. Himachal Pradesh
. NMW 18569,
NMW18570:1–2, “Simla and Kulu (Himalaya)”, now Shimla and Kullu (Syntypes of Zamenis himalayanus Steindachner,
1867).
Sikkim. FMNH 15827, “Mangpu, Sikkim”. West Bengal. NMW 22383:2, NMW 22383:5, Darjeeling.
Amphiesma sieboldii (n = 23). Nepal. BMNH 1913.5.22.1, “Maikola Valley, E. Nepal”, now Mai Kola; CAS 90690, “Nepal:
above Deppur (Elev. 5500 Ft)”; FMNH 109762, Amp Pipal, 4000'; FMNH 131966, Chapagaon, Kathmandu Valley;
FMNH 131967, Kathmandu Valley; FMNH 190856, Arun Valley, at Num Bridge across Arun River; FMNH 204499,
above Num, 6400' in forest area; FMNH 204500–504, no precise localities; MHNG 1355.72–73, Astam, near Hyangcha,
1600m; MNHN 2003.3614, east of Mounasko Pass, between Surkie Pass and Chheskam, Eastern Region, 2400 m;
ZMB 4551, “Himalaya”; ZMB 10231, Sikkim. - India. West Benga
l. CAS-SU 15973, Darjeeling; NMW 22383:1, NMW
22383:3–4, Darjeeling. - People’s Republic of China. Xizang Autonomous Region (Tibet)
. CAS 177474, elev.
2,000–2,100 m, between Chinese check point at Zhangmu (Khasa) (28°07'N-85°59'E) and the Nepal border on the Lhasa-
Kathmandu Rd., Xigaze Prefecture; CAS 177672–177673, elev. 2,300–2,500 m, between Chinese check point at Zhangmu
(Khasa) (28°07'N-85°59'E) and the Nepal border on the Lhasa-Kathmandu Rd., Xigaze Prefecture.
