Genus Amphiesma Dumeril, Bibron & Dumeril, 1854

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ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2013 Magnolia Press

Zootaxa 3694 (4): 301–335

www.mapress.com/zootaxa/Article

http://dx.doi.org/10.11646/zootaxa.3694.4.1

http://zoobank.org/urn:lsid:zoobank.org:pub:6F2F58C1-3927-4E66-B09C-AFAF4B4A2C71

On some taxonomically confused species of the genus Amphiesma Duméril,

Bibron & Duméril, 1854 related to Amphiesma khasiense

(Boulenger, 1890) (Squamata, Natricidae)

PATRICK DAVID1, GERNOT VOGEL2 & JOHAN VAN ROOIJEN3

1Reptiles & Amphibiens, UMR 7205 OSEB, Département Systématique et Evolution, CP 30, Muséum National d’Histoire Naturelle, 57

rue Cuvier, 75231 Paris Cedex 05, France. E-mail: [email protected] for Southeast Asian Herpetology, Im Sand 3, D-69115 Heidelberg, Germany. E-mail: [email protected] 313, 2272 EH Voorburg, The Netherlands. E-mail: [email protected]

Table of contents

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302

Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303

Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 304

Amphiesma khasiense (Boulenger, 1890) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 306

Amphiesma boulengeri (Gressitt, 1937) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312

Amphiesma inas (Laidlaw, 1901) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 318

Amphiesma kerinciense David & Das, 2003. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 321

Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 324

Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 328

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329

Literature cited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329

Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 335

Abstract

Three species of the genus Amphiesma Duméril, Bibron & Duméril, 1854 have long been confused in the literature, with

each other and with other species of the genus. Amphiesma khasiense (Boulenger, 1890) has been considered to inhabit a

large geographical region, extending from north-eastern India, east to Vietnam and southern Thailand. Amphiesma bou-

lengeri (Gressitt, 1937) has been regarded as a species endemic to south-eastern China. Amphiesma inas (Laidlaw, 1901)

has been recorded from West Malaysia, Thailand and Indonesia (Sumatra). A multivariate analysis of morphometric and

meristic characters shows that these three species can be separated by combinations of characters in the scalation and pat-

tern, the most obvious being the structure of the postocular streak. On the basis of our analysis and after comparison with

name-bearing type specimens, Amphiesma khasiense is restricted to north-eastern India, Myanmar, western Yunnan Prov-

ince of China, northern Laos and northern and western Thailand. Other populations from south-eastern China, Vietnam,

other parts of Laos, Cambodia and central Thailand, which have been recorded in the literature as A. khasiense, A. johannis

or Amphiesma modestum (Günther, 1875), should be referred to Amphiesma boulengeri. Amphiesma inas (Laidlaw, 1901)

is a valid species endemic to mountain ranges of southern Peninsular Thailand and West Malaysia. The mention of Am-

phiesma inas in Sumatra is erroneous, being based on the second known specimen of Amphiesma kerinciense David &

Das, 2003, which is here redescribed. A key to species of the Amphiesma khasiense group and other species sharing a

greyish-brown background without conspicuous dark and pale stripes, is provided.

Key words: Serpentes, Natricidae, Amphiesma, Amphiesma khasiense, Amphiesma boulengeri, Amphiesma inas, Am-

phiesma kerinciense, Indochina, taxonomy

Accepted by Z. Nagy: 4 Jul. 2013; published: 6 Aug. 2013 301


Introduction

On the basis of morphological characters, especially the hemipenial morphology and dentition, Malnate (1960)

split the speciose genus Natrix Laurenti, 1768, into several genera, including Amphiesma Duméril, Bibron &

Duméril, 1854. This latter genus was itself divided into several informal phenetic groups of species. Malnate

recognized two informal, unnamed groups dealing with species of interest here. The first one, which we name here

the “Amphiesma modestum species group”, included A. modestum (Günther, 1875), A. atemporale (Bourret, 1934),

A. boulengeri (Gressitt, 1937), A. deschauenseei (Taylor, 1934), A. groundwateri (Smith, 1922), A. inas (Laidlaw,

1901), A. johannis (Boulenger, 1908) and A. xenura (Wall, 1907). A second group, with morphological and

hemipenial characters close to those of the first group, and named here the “Amphiesma khasiense species group”,

includes 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. Malnate (1962: 252) eventually placed A. craspedogaster, A.

popei, A. pryeri, A. sauteri and A. vibakari into two distinct species groups, formally named as the Amphiesma

vibakari species group (for A. vibakari, A. popei and A. sauteri) and the Amphiesma pryeri species group (for A.

pryeri, A. ishigakiensis and A. craspedogaster). However, Malnate (1960) had pointed out that Amphiesma

parallelum and 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 this character as well as other morphological

characters which will be discussed elsewhere, we consider Amphiesma parallelum and related species to constitute

a different species group. Due to their morphological characters, we do the same for species related to Amphiesma

modestum, i. e. A. deschauenseei, A. groundwateri, and A. modestum. The relationships of A. xenura are unclear.

Furthermore, A. venningi, with 17 dorsal scale rows at midbody, shares most external characters with the group of

A. modestum and other undescribed species (David et al., in prep.). This leaves only seven species in the artificial

“Amphiesma khasiense species-group” of interest here: A. khasiense, A. atemporale, A. boulengeri, A. inas, A.

johannis and two recently described species, Amphiesma kerinciense David & Das, 2003 and Amphiesma

leucomystax David, Bain, Nguyen, Orlov, Vogel, Vu & Ziegler, 2007. These two latter species are referred to this

group by their habitus, scalation and pattern (David & Das 2003; David et al. 2007).

According to Malnate (1960) and our data, the expanded “A. khasiensis species group” can defined by (1)

hemipenes single, (2) sulcus spermaticus single, (3) maxillary teeth progressively increasing in size posteriorly, (4)

internasal scales broadly truncated anteriorly, (5) nostrils lateral, (6) 17 (A. atemporale) or 19 dorsal scale rows

(other species) at midbody, and (7) a dorsal pattern made of aligned dots or blotches on more or less faint

dorsolateral stripes.

Amphiesma khasiense was described from the the Khasi Hills in north-east India. Smith (1943) also recorded it

from northern Burma and northern Vietnam. Subsequently, this species has been widely cited from Yunnan

Province in China (Zhao & Adler 1993), from Vietnam and other countries of the Indochinese Peninsula (see, for

example, Smith 1943; Nguyen et al. 2009), and eventually from northern Thailand (Chanhome et al. 2001;

Pauwels et al. 2009). From another geographical direction, Gressitt (1937) described Natrix boulengeri from

Guangdong Province in south-eastern China. Both species are morphologically rather similar, especially in their

general dorsal pattern. A casual examination of preserved specimens and of pictures published in the literature

showed that several, if not many authors since Smith (1943) have confused A. khasiense, A. boulengeri, A. inas and

even A. modestum.

In an attempt to precise the definitions of species of the A. khasiense species group present in the Indochinese

Peninsula and the Indo-Malayan Region, we examined a total of 142 specimens covering all species of the

Amphiesma khasiense species group as listed above. Although species such as A. atemporale, A. johannis, A.

kerinciense and A. leucomystax are easily diagnosed, our material revealed morphological differences in scalation,

morphometry and dorsal and head patterns which were not in agreement with the diagnoses and ranges given to A.

khasiense sensu stricto of the Indo-Himalayan Region, “A. khasiense” auctorum of the Indochinese Peninsula, A.

boulengeri and A. inas in the literature. In this paper, we present morphological analyses of this material, especially

multivariate analysis of morphometric and meristic characters which lead to redefinitions of these species.

DAVID ET AL.302 · Zootaxa 3694 (4) © 2013 Magnolia Press


Material and methods

The revision is based on external morphological characters regarded as taxonomically significant in the genus

Amphiesma as defined by Malnate (1960), Malnate & Underwood (1988), Ziegler & Le (2006a), David et al.

(2007) and David & Vogel (2010). A total of 142 specimens of A. khasiense, A. boulengeri, A. inas, and A.

kerinciense were examined, of which 130 were used for multivariate analyses. These specimens are listed under the

respective species accounts. Additional specimens of other Amphiesma species are listed in the Appendix.

Morphological data. Maxillary teeth were counted by dissecting the right maxilla of one specimen and, for

other specimens, in removing the exterior gum surfaces of the jaw in situ. Dentitional data of other species of

Amphiesma were obtained in the same way (tooth sockets were included in the counts in cases of tooth loss).

Measurements, except body and tail lengths, were taken with a slide-calliper to the nearest 0.1 mm; all

measures on body were measured to the nearest millimetre. The number of ventral scales is counted according to

Dowling (1951). The numbers of dorsal scale rows are 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. The terminal scute is not included in the number of subcaudals. Values for symmetric head characters

are given in left / right order.

Abbreviations. Measures and ratios. DEYEH: horizontal diameter of the eye. DEYEV—vertical diameter of

the eye. DLIP—distance between the lower edge of the lip and the lower margin of the eye. HL—head length.

LFRO—length of frontal. LPARIET—length of parietal. LPREFRO—length of prefrontal. SnLERO—snout

length between eye and anterior margin of rostral. SVL—snout-vent length. TaL—tail length. TL—total

length.TaL / TL—ratio tail length / total length. WFRO—Width of frontal. WINASP—width of internasals

(posteriorly).TEETH—number of maxillary teeth.

Meristical characters. DSR—dorsal scale rows. IL—infralabials. K1DSRM—keeling of scales of 1st dorsal

scales at midbody. K1DSRR—keeling of scales of 1st dorsal scales on the posterior part of the body. KMSR—

keeling of dorsal scales at midbody. POSOCMED—mean number of postoculars. SC—subcaudals. SL—

supralabials. VEN—ventrals. REDUC1—position (in VEN) of the first reduction of dorsal scale rows. REDUC2—

position (in VEN) of the second reduction of dorsal scale rows. SC4—extension, expressed in number of

subcaudals, of 4 scale rows around the tail. SC6—extension, expressed in number of subcaudals, of 6 scale rows

around the tail.

Pattern characters. DORSPATT: General dorsal pattern: 1: uniform; 2: variegated; 3: strong fasciatures.

NAPE—Pattern of the nape: 0: none; 1: isolated light blotches; 2: narrow white (cream) continuous stripe; 3: wide

(cream) continuous stripe; 4: wide stripe continuing dorsolateral stripe; 5: broad white stripe (leucomystax).

PatPoStr—Pattern of the postocular streak: 1: blotches centered, rounded; 2: elongated blotches, not in contact; 3:

narrow nearly continuous stripe; 4: wide continuous stripe; 5: low, triangular blotches. POSITDSL—position of the

dorsolateral stripe (in number of scale rows): 1: 5-6; 2: 5-7; 3: 6-7. PosPoStr—position of the postocular streak: 1:

blotches centered in SL 7-8-9; 2: blotches on upper SL7, middle SL 8 and 9; 3: middle of 7, lower 8 and 9; 4: low

7, 8 and 9. STRIPE—0: none; 1: faint stripe with aligned dots; 2: distinct, continuous stripe; 3: vertical or divided

spots. VENTPATT—0: none; 1: faint dark blotch on ventral tips; 2: double blotch on tips.

Multivariate analyses. For the analysis of geographic variation within A. khasiense, A. boulengeri and A.

inas, specimens were grouped into 14 OUT’s detailed in Table 1.

Subsequently, characters were selected that exhibited significant geographic variation. Numerical variables

pertaining to morphology were analyzed with (M-)ANCOVA (e.g. Maxwell & Delaney, 1990) using OTU (Table

1) and sex as factors and SVL as covariate. Categorical variables (pertaining to coloration or the degree of keeling

of the dorsal scales) were analyzed with a χ2- test. Characters which exhibited significant geographic variation at

the (bonferroni-corrected) 0.001-level were selected for further multivariate analysis. However, characters with too

many missing values were omitted.

For multivariate analyses, morphometric variables were adjusted to a common SVL of 32.5 cm to correct for

potential ontogenetic variation between the samples of the species (e.g. Thorpe 1975, 1983; How et al. 1996; Turan

1999). The following allometric equation was applied: Xadj

= X - β(SVL - SVLmean

) where Xadj

is the adjusted value

of the morphometric variable; X is the original value; SVL is the snout-vent length; SVLmean is the overall mean

snout-vent length; β is the coefficient of the linear regression of X against SVL. Linearity of the relation between

SVL and each morphometric variable was checked by visual inspection of the scatter plot.

Zootaxa 3694 (4) © 2013 Magnolia Press · 303REVISION OF THE GROUP OF AMPHIESMA KHASIENSE


TABLE 1. OTU’s and corresponding sample-sizes. The specific names used in this table are based on those most

frequently found in the literature before the present work.

Nonlinear Principal Component Analysis (PCA; Shen & Lai 1998; Cramer 2003) was used to reduce the

dimensionality of the data set. Variables pertaining to coloration were specified as nominal, variables pertaining to

the degree of keeling of the dorsal scales as ordinal and all others as numerical. In order to visualize possible

discrete phenotypic transitions, the object scores corresponding with the first principal component were plotted, for

males and females separately, against longitude and latitude in at three-dimensional plot. The objective of this

analysis was to enable differentiation between clinal variation and phonetic discontinuities, the latter being

evidence for lineageseparation (e.g. Lenk & Wüster 1999; De Queiroz 2007).

All statistical analyses were carried out with the software SPSS (2006; SPSS for Windows. Release 14.0.2.

Chicago: SPSS Inc.).

Museum abbreviations: AMNH—The American Museum of Natural History, New York, USA. BMNH—

Natural History Museum, London, UK. CAS—California Academy of Sciences, San Francisco, USA. CIB—

Chengdu Institute of Biology, Chengdu, People’s Republic of China. CTNRC—Center for Thai National Reference

Collections, Bangkok, Thailand. FMNH—Field Museum of Natural History, Chicago, USA. IEBR—Institute for

Ecology and Biological Research, Hanoi, Vietnam. KFBG—Kadoorie Farm and Botanic Garden, Hong Kong

SAR, People’s Republic of China. KZM—Korat Zoo Museum, Korat. LSUHC—La Sierra University

Herpetological Collection, Riverside, California, USA. MNHN: Muséum National d’Histoire Naturelle, Paris,

France. MVZ—Museum of Vertebrate Zoology, Berkeley, California, USA. MZB—Bogor Zoological Museum,

Bogor, Indonesia. NCSM—North Carolina State Museum of Natural Sciences. PSGV—Gernot Vogel’s private

collection, Heidelberg, Germany. ROM: Royal Ontario Musem, Toronto, Canada. QSMI: Queen Saovabha

Memorial Institute, Thai Red Cross, Bangkok, Thailand. SMNH: Shanghai Museum of Natural History, Shanghai,

People’s Republic of China. ZISP—Zoological Institute, Russian Academy of Science, St. Petersburg, Russia.

ZMB: Zoologisches Museum für Naturkunde der Humboldt-Universität zu Berlin, Berlin, Germany. ZRC—

Zoological Reference Collection, Raffles Museum of Biodiversity Research, National University of Singapore,

Singapore. ZSI—Zoological Survey of India, Kolkata, India.

Results

Table 2 shows the selected characters and their correlations with the first principal component in the multivariate

analysis run on populations referred to as A. khasiense, “A. khasiense” auctorum of the Indochinese Peninsula, A.

boulengeri and A. inas.

OTU Sample size

1 khasiense Myanmar / India 25

2 khasiense China (Yunnan) 2

3 khasiense North Thailand 4

4 khasiense Laos 2

5 inas South Thailand and West Malaysia 8

6 boulengeri South-east China 7

7 boulengeri Northern Vietnam 2

8 cf. khasiense South-east China 3

9 cf. khasiense Northern Vietnam 34

10 cf. khasiense Central Laos 2

11 cf. khasiense CentralVietnam 17

12 cf. khasiense Southern Vietnam 8

13 cf. khasiense Cambodia 14

14 cf. khasiense East Thailand 2

TOTAL 130



TABLE 2. Correlations between original variables and first principal component.

A plot of the first principal component against longitude and latitude is shown in Fig. 1.

Figure 1 shows clear, sharp phenotypic transitions between A. khasiense, A. boulengeri and A. inas. Specimens

referred to “A. khasiense” of the Indochinese Peninsula are shown to not be distinguishable from A. boulengeri.

Thus, the analyses supports the existence of only three taxa recognized at the species level, A. khasiense, A.

boulengeri and A. inas. The specific nomen khasiensis should be restricted to populations of OTU 1–4 as defined in

Table 1. On another side, Amphiesma inas contains only specimens from Peninsular Thailand and West Malaysia.

Populations from OTU 6–13 should all be referred to Amphiesma boulengeri. However, visual inspection of the

data suggested morphological divergence of the Cambodian population of A. boulengeri (OTU 13) from the other

populations (OTU’s 6-12). A MANCOVA was carried out to establish in what morphological characters OTU 13

differs significantly from OTU’s 6-12. These characters (VEN, SC, REDUC1, REDUC2, LPREFRO, WFRO,

WINASP, Sc6, Sc4) were subsequently included in a PCA and the resulting first PC was plotted against longitude

and latitude (Fig. 2).

Although Fig. 2 clearly shows morphological divergence of the Cambodian population, it does not rule out the

possibility that this is the result of steep clinal variation. Therefore, we conservatively consider the Cambodian

population to be conspecific with the other populations of A. boulengeri. In conclusion, our results do not provide

clear evidence that more taxons are involved, especially under the name A. boulengeri. A. kerinciense, endemic to

Sumatra and morphologically relatively distinct from the three other species, was not considered in these analyses.

The recognized are redefined below. If we provide complete synonymies and chresonymies, i.e. the various

combinations under which the currently valid species has been known in the literature, by far the citations of the

literature cannot be considered to be exhaustive.

Character

VEN 0.15 0.10

POSOCMED 0.16 0.49

KMSR 0.81 0.03

K1DSRM 0.91 0.20

K1DSRR 0.92 0.15

PosPoStr -0.89 -0.21

PatPoStr -0.87 -0.21

NAPE -0.89 -0.09

DORSPATT 0.18 0.39

POSITDSL -0.63 0.01

STRIPE -0.18 -0.37

VENTPATT -0.31 -0.34

SnLERO -0.60 0.33

DEYEH -0.10 0.73

DEYEV -0.19 0.86

DLIP -0.57 0.48

LPARIET -0.33 0.29

LFRO -0.31 0.62

WFRO -0.69 0.16

LPREFRO -0.40 0.31

TEETH -0.70 -0.15



FIGURE 1. Object scores of first principal component along the geographic coordinates: circles: A. khasiense, males; triangles pointed upwards: A. khasiense, females; triangles pointed downwards: “A. cf. khasiense” of the Indochinese Peninsula, males; triangles pointed to the right: “A. cf. khasiense” of the Indochinese Peninsula, females; triangles pointed to the left: A. boulengeri, males; pentagons: A. boulengeri, females; squares: A. inas, males; diamonds: A. inas, females.

Amphiesma khasiense (Boulenger, 1890)

(Fig. 3, 4D–F, 5)

Tropidonotus khasiensis Boulenger, 1890: 344.—Type locality. “Khasi Hills”, in the State of Meghalaya, India.—Syntypes (4). BMNH 1946.1.12.80 (adult female) and 1946.1.12.81–82 (adult males; all formerly BM 70.11.30.33), collected by T. C. Jerdon; BMNH 1946.13.45 (juvenile; formerly 53.8.12.44); collected by Sir. J. Hooker.

Tropidonotus khasiensis.—Boulenger 1893a: 322; 1893b: 223, Pl. 13: Fig. 3 - Fea 1897a: 475; 1897b: 95 - Annandale 1905: 210; 1912: 49 & 53 - Wall 1908: 316, 317: Fig. 2 - Werner 1929: 24.

Natrix khasiensis.—Wall 1923: 601; 1926: 559 - Mell 1929: 148 (in part) - Bourret 1936a: 103, 113 & 132 (in part); 1936b: 69, Fig. 28 (in part for the text; see under Amphiesma boulengeri for the figure) - Smith 1940: 483; 1943: 289.—Deuve 1970: 84 & 88 (in part). - Anonymous 1977: 63 & 87 - Djao et al. 1977: 69 - Yang et al. 1978: 67; 1983: 43.

Amphiesma khasiensis.—Malnate 1960: 50, 52 & 57 - Tian et al. 1986: 116 & 149—Yang & Inger 1986: 7—Zhao 1986: 239—Zhao et al. 1986: 201; 1998: 52 & 62: Fig. 18—- Hu et al. 1987: 126—Welch 1988: 31—Zhao & Adler 1993: 226—Das 1996: 53—Zhao & Yang 1997: 201 & 207—Mathew 1998: 132—Sharma 1998: 94—Orlov et al. 2000: 71 (in part)—Ji 2002: 176 & 177—Hallermann et al. 2002: 150—Sharma 2003: 132 & 133; 2007: 206 & 209 (in part)—Zhao 2006: (I)164, (II)85: Fig. 48—Yang & Rao 2008: 258—Li et al. 2010: 166 & 209: Fig. 3-14—Luo et al. 2010: 74—Murthy 2010: 33.

Amphiesma khasiense.—Zhao et al. 2000a: 232; 200b: 205—Chanhome et al. 2001: 53 & 57.—Iskandar & Colijn 2001: 96—He & Zhou 2002: 167—Das 2003: 473; 2010: 150: Pl. 67: Fig. 9, 332 (in part for the text; at the exception of of the mentions of Thailand [Loei Province] and “northern Vietnam”)—Whitaker & Captain 2004: 25, 218 & 219, Figure—David et al. 2007: 54 & 55—Sen & Mathew 2008: 140, 159, Fig. 160–161—Ahmed et al. 2009: 19—Pauwels et al. 2009: 75—Teynié & David 2010: 228, 284 & 308.



Natrix gilhodesi Wall, 1925a: 587, Plate: Fig. 2, 2a–c. – Type locality. “Huton, Bhamo District (30 miles north-east of Bhamo; circa 4,500 feet; Lat. Circa 97° .33; Long. Circa 24° .24)”, now Hutung (24°15'N-97°31'E; 1,715 m a.s.l.), Kachin State, Myanmar. – Holotype. BMNH 1946.1.13.62 (adult male; formerly BM 1925.4.2.8); collected by Father Gilhodes and deposited by F. Wall (see below). – Status. A junior subjective synonym of Tropidonotus khasiensis Boulenger, 1890; synonymized by Wall (1926: 559).

Natrix gilhodesi.—Wall 1925b: 808.Natrix kashiensis (lapsus calami pro Natrix khasiensis).—Mell 1931b: 225 & 238.Amphiesma modesta (nec Tropidonotus modestus Günther, 1875, a distinct species).—Cox et al. 1998: 45—Das 2002: 18.Paranatrix modesta (nec Tropidonotus modestus Günther, 1875, a distinct species).—Mahendra 1984: 247 (see below).Amphiesma inas (nec Natrix inas Laidlaw, 1901, a distinct species).—Chan-ard et al. 1999: 153 (top; specimen from Phu

Luang, Loei Province, Thailand)—Nabhitabhata et al. 2004: 124 (in part: mention of Phu Luang, Loei Province)—Das 2010: 332 (only for mentions of Thailand: Loei and Nakhon Ratchasima Provinces).

FIGURE 2. Object scores of first principal component along the geographic coordinates: circles: A. boulengeri, Cambodia, males; triangles pointed upwards: A. boulengeri, Cambodia, females; triangles pointed downwards: A. boulengeri, males; triangles pointed to the right: A. boulengeri, females.

Material examined (35 specimens). – India. State of Meghalaya. BMNH 1946.1.12.80–1946.1.12.82, BMNH

1946.1.13.45, Khasi Hills. State of Arunachal Pradesh. P 377–379, Dihang Dibang Biosphere Reserve, District of

Dibang Valley; ZSI 23926, Changlang. State of Nagaland. KSC 140, Kohima. – Myanmar. Chin State. CAS

220023, Nat Ma Taung National Park (21°22'20.1N 93°58'34.6E), Min Dat Township, Min Dat District. Kachin

State. BMNH 1946.1.13.62 (holotype of Natrix gilhodesi Wall, 1925), BMNH 1946.1.13.63, BMNH 1925.4.2.10–

15, BMNH 1925.4.2.15A, “Huton, Bhamo District (30 miles north-east of Bhamo; circa 4,500 feet; Lat. Circa 97° .33; Long. Circa 24° .24)”, now Hutung; BMNH 1974.884, Nawng Hkai, near Putao; CAS 221504, between Babaw

and Rabaw (27°26'28.4N 97°55'06.3E), Naung Mon Township, Putao District; CAS 221525, Rabaw (27°26'28.4N

97°55'07.5E), Naung Mon Township, Putao District; CAS 221543, Rabaw (27°26'14.9N 97°55'21.1E), Naung



Mon Township, Putao District; CAS 224654, Nagmung (27°30'18.8N 97°48'33.9E), Nagmung Township, Putao

District; CAS 224694, Nagmung (27°29'49.6N 97°49'06.9E), Nagmung Township, Putao District. Kayah State.

MNHN 1893.0399, Monts Karen. – People’s Republic of China. Yunnan Province. CIB 2000I0009, Ruili; CIB

Xi0089, Xichuangbanna. – Laos. Phongsaly Province. MNHN 2004.0248, Long Nai Tai. – Thailand. Chiang Rai

Province. KZM 001, QSMI 542, near Ban Pa Miang Mae Hang, Moo. 7, Pagnew Subdistrict, Wieng Pa Pao

District. Chiang Mai Province. CTNRC 980504, Doi Inthanon National Park; FMNH 251780–251781, Chiang

Mai. Loei Province. QSMI 0273, Phu Luang.

Taxonomic comments. Amphiesma khasiense is here redefined as an Indo-Himalayan species which enters the

mountain ranges of North Thailand and northern Laos. In the literature, this species has often been confused with

Amphiesma boulengeri as defined here, and, more rarely, with A. inas. These confusions explain the numerous

citerations in part in the chresonymy given above. This species is monotypic.

We have examined the holotype of Natrix gilhodesi Wall, 1925 and we concur with Smith (1943) in regarding

this taxon as a junior synonym of Amphiesma khasiense. Furthermore, Wall (1925a: 587) stated he had six

specimens of Natrix gilhodesi at hand but that one was selected as the type. This specimen is registered as BMNH

1946.1.13.62. According to the catalogue of the Natural History Museum of London, other specimens of the

original series are BMNH 1946.1.13.63 and BMNH 1925.4.2.10–15, BMNH 1925.4.2.15A, so a total of eight

specimens.

Mahendra (1984: 244) erected the genus Paranatrix (type species: Tropidonotus modestus Günther, 1875, now

Amphiesma modestum, by original designation) for six species of the genus Amphiesma present in India.

Furthermore, Mahendra (1984: 247) synonymized A. khasiense with A. modestum. This synonymy is unjustified as

these species are morphologically distinct as shown above. However, the generic nomen Paranatrix is an available

name if the genus Amphiesma has to be split.

Diagnosis. A species of the genus Amphiesma characterized by the combination of (1) body slender in males

and females; (2) nostrils directed laterally; (3) 21–26 maxillary teeth, gradually enlarged, the last 2 or 3 moderately

enlarged; (4) 19 dorsal scale rows at midbody, strongly keeled at the exception of scales the 1st dorsal scale row,

smooth or weakly keeled on the anterior half of the body, weakly or distinctly keeled posteriorly; (5) dorsal pattern

made of a series of reddish-brown or rusty dots or small blotches, usually aligned on a faint dorsolateral stripe; (6)

each of the three or four posterior supralabials with a large, white, cream or ivory, rounded, subrectangular,

triangular or vertically oval blotch, or, rarely, nearly entirely cream or ivory with dark edges; (7) pattern of the neck

usually made of isolated rounded blotches, sometimes partly connected by a narrow pale line; (8) venter ivory or

cream, with the outer quarter of each ventral dark brown, separated from the dark dorsal colour by a narrow pale

line; (9) eye large, 1.8–2.8 times the distance between the lower margins of eye and of lip; (10) 141–155 VEN, 87–

111 SC; (11) internasals abruptly truncated anteriorly; (12) 9 (rarely 8) supralabials; (13) 1 anterior (rarely 2)

temporals.

A. khasiense, A. boulengeri and A. inas are morphologically similar. Characters of use for separating these taxa

are given below in the Discussion.

Variation (based on 35 examined specimens and Smith 1943). Body cylindrical, elongate, rather slender in

males, slightly more robust in old females; head short, oval, distinct from the thick neck, flattened; snout short,

blunt, amounting for 24.5–30.4 % of HL or 1.2–1.4 times as long as diameter of eye; nostrils large, rounded,

directed laterally and piercing in the middle of the nasal; eye large, 1.8–2.8 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 known is 673 mm according to Wall (1923b: 601); the largest specimen seen by us is

604 mm (SVL 412 mm; TaL 192 mm; specimen BMNH 1946.1.12.80; female). The longest known male is 562

mm long according to Wall (1926); the longest male seen by us is 549 mm long (SVL 368 mm, TaL 181 mm long;

BMNH 1925.4.2.11). Ratio TaL / TL: 0.296–0.346, without clear sexual dimorphism (see below).

Dentition. Maxillary teeth 21–26, gradually enlarged, the last 2 or 3 teeth distinctly enlarged.

Body scalation. DSR: 19–19–17 scale rows; dorsal scales rhomboedric, moderately to strongly keeled on all

rows but 1st DSR, normal in shape or notched posteriorly; scales of 1st DSR weakly keeled on the anterior half of

the body, rarely smooth, posteriorly weakly keeled in juvenile specimens but moderately to strongly keeled in

adults.

Scale row reductions: first reduction (19→17) at VEN 89–96 (x = 92.5; s = 2.5); second reduction (17→15) at

VEN 92–99 (x = 95.5; s = 2.1).



VEN: 141–155 (plus 1 or 2 preventrals); SC: 87–111, all paired, with a sexual dimorphism; ratio VEN / SC:

1.37–1.69; anal plate divided.

Position of the reduction from 8 to 6 scale rows around the tail: 8th–24th 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:

0.88–1.31 with a weak sexual dimorphism.

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 0.8–1.1 times as long as wide, abruptly truncated anteriorly with anterior

margin about 0.55–0.75 times the width of the posterior margin; 2 wide prefrontals, 1.1–1.5 times as long as

internasals; frontal shield-like with apex directed posteriorly, moderately enlarged, 1.3–1.5 times longer than wide,

2.0–2.5 (usually 2.1–2.2) times as long as prefrontal; 1 supraocular on each side, longer than wide, about as wide as

internasals; two large parietals, 1.3–1.6 times longer than frontal; 1 / 1 subrectangular loreal scale, 0.8–1.0 times as

high as long, in broad contact with the nasal; 9 (exceptionally 8) SL (9 / 9 in 37 cases out of 39, 8 / 8 in 1 case, 8 /

9 in 1 case), 3rd–9th longer than high; 1st and 2nd SL small and short, in contact with nasal; 2nd and 3rd SL or more

rarely 2nd–4th SL in contact with the loreal; 4th–6th SL (or exceptionally 3rd–5th SL in 3 out of 70 occurrences)

entering orbit; 7th and 8th SL largest; 1 or 2 preoculars on each side (2 preoculars in only 18 / 70 occurrences); 3

(exceptionally 2 in 1 / 70 occurrences) small postoculars; 1 anterior temporal (2 in 4 / 70 occurrences), narrow and

elongate, with complete temporal formulas as 1+1+2, 1+2+2, more rarely 2+1+2 or 2+2+2; usually 10 / 10

infralabials (rarely 10 / 11, 9 / 9 in only 1 out of 35 specimens), first pair in contact each with other, 1st–5th IL in

contact with anterior chin shields, 6th IL largest.

Coloration and pattern in alcohol. The dorsal and lateral surfaces of body are ochre-brown, greyish-brown,

dark gray, dark grayish-brown or dark chestnut brown, often distinctly darker, i.e. dark grayish-brown or blackish-

brown, on the five upper scale rows of the back, throughout more or less heavily variegated with much darker

pigmentation; back and each side with 2 or 3 series of small, blackish-brown spots, subrectangular or elongated,

more or less well defined and scattered in alternance each with the other; on each side, a faint, ochre or reddish-

brown dorsolateral stripe extending from the neck on upper half of 5th and the whole of 6th dorsal scale rows,

sometimes also on lower part of scales of the 7th row is often present although often not visible after midbody; on

each dorsolateral stripe, a series of small, irregular pale, dark-edged spots, cream, yellow, ochre or rusty brown

(coral, orange or red in life), covering only one scale and arranged every 1 to 3 or 4 dorsal scales, conspicuous on

the anterior part of the body, progressively fainter and often not visible on the second third of body; some paler

dashes may be present between the dorsolateral spots on the anterior part of the body; 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 15th–20th ventral. The

tail is as the body, with minute dorsolateral cream or yellow (coral, orange or red in life) spots, vanishing

progressively; posterior half of tail uniformly dark, usually variegated with irregular and faint darker pigmentation.

The head is ochre-brown, greyish-brown, dark gray or dark grayish-brown, paler than the background colour

of the body, often paler 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; a short cream streak just

behind the suture of parietals present or absent; a pattern made of twin pale spots, one on each parietal, present or

absent; 1st-5th, sometimes 1st-6th anterior supralabials white, cream or pale yellow (same in life), narrowly edged

with dark brown or black; 1st supralabial sometimes darker than other anterior supralabials; 6th to 9th or 7th to 9th

supralabials with a large, conspicuous blotch in their centre, pure white or cream and broadly edged with blackish-

brown, rounded or elongate (i.e. subrectangular, slightly horizontally ovale or rarely triangular), straight or slightly

oblique, more rounded in 9th SL, not in contact with each other and contrasting sharply with the dark colour of the

head; exceptionally the posterior supralabials are nearly entirely white, cream or ivory with dark edges (see below);

no postocular streak; on each side of the neck, from the corner of the mouth behind 9th supralabial, an oblique series

of small, isolated, rounded or elongate white or cream blotches (same colour in life), usually not connected with

each other or sometimes partly connected by a narrow pale line, usually not reaching the upper part of the neck;

these blotches may even be reduced to a few pale spots and do not constitute a continuous stripe. The throat and

chin are cream, ivory or very pale yellow (pale pink or reddish-gray in life); infralabials and mental more or less

heavily dotted or speckled with small, irregular dark brown spots; sometimes minute dots on the preventral area.

The pupil is black, the iris is dark red in life.



The venter is uniformly ivory, cream, pale yellow or pale ochre (bright pink or reddish-gray in life); about a

quarter of the outer part of each scale with a well-defined, sharp dark brown or blackish-brown area, separated

from the dark similar colour covering the ventral tip by a very narrow, cream or ivory line or streak at the base of

the tip, crossing the scale or limited to a spot at the posterior edge of the ventral, 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

heavily speckled with minute dark brown dots, nearly entirely dark near its tip.

The pattern of the posterior supralabials, i.e. a rounded or elongate, isolate white, cream or ivory central blotch

is diagnostic of this species. However, exceptionally, the posterial supralabials may be nearly entirely pale, with

only dark edges. We have met only two such specimens, MNHN 1893.0399 (Karen Mts, Myanmar; identified as A.

khasiense by E. Malnate according to the catalogue of the MNHN) and MNHN 2004.0248 (Long Nai Tai,

Phongsaly, Laos; depicted in Teynié & David 2010: 229) respectively. These specimens may easily be identified as

Amphiesma modestum but the keeled 1st dorsal scale row, pattern of the neck and the well-defined lateral blotches

of the venter is different and diagnostic of A. khasiense.

Hemipenis. In situ, it is short and stout, undivided but with two short extensions at its tip, reaching in situ the

8th or 9th SC; sulcus spermaticus simple, S-shaped, reaching the top of the hemipenis, with prominent lips;

hemipenis entirely covered with short but dense spines at the exception of the base which is covered with large

calyces; lips of the sulcus also covered with short, thick spines; both extensions densely covered with rather

numerous thin spines; 2 or 3 large, thick spines on the internal side of the organ at 90° of the sulcus.

Sexual dimorphism. It is expressed only in the difference in the number of subcaudals, with males: 96–111

(x = 103.3, s = 4.2) vs. females: 87–97 (x = 93.8, s = 3.5). Another weaker difference appears in the position

(counted in number of subcaudals) of the reduction from 6 to 4 scale rows around the tail, with males: 15th–22nd SC

(x = 18.7, s = 2.8) vs. females: 8th–21st SC (x = 13.4, s = 4.8).

Distribution. According to authors cited above in the chresonymy and our material, the distribution of A.

khasiensis is currently known as follows: India. Known only from north-east India. State of Meghalaya: Khasi

Hills and Garo Hills. State of Arunachal Pradesh: Changlang; Dibang. State of Nagaland: Kohima District; Wokha

District.—Myanmar. Kachin State. Regions of Bhamo, Hutung, Putao and Sumka Uma (Mahtum, Sumka Uma).

Chin State. Nat Ma Taung National Park, Min Dat District. Kayah State. Monts Karen.—People’s Republic of

China. Yunnan Province. Xichuangbanna, in extreme southern Yunnan, and Ruili. Xizang Province. South of

Motuo.—Laos. Recorded only from the north of the country. Phongsaly Province. Long Nai.—Thailand. Known

only from the mountainous regions of the northern and western parts of the country. Chiang Rai Province. Near

Ban Pa Miang Mae Hang, Pagnew, Wieng Pa Pao District. Chiang Mai Province. Chiang Mai. Loei Province. Phu

Luang. Tak Province. Umphang; photographed specimen (Fig. 5 top left). See Pauwels et al. (2009) for a detailed

discussion on the distribution of this species in Thailand.

Records from Vietnam, Cambodia and other parts of Thailand should be referred to Amphiesma boulengeri or

A. inas (see below).

Biology. This rather uncommon species inhabits wet primary submontane and montane forests and wet

secondary forests between about 600 and 1,400 m a.s.l. It is usually associated with the forest floor near the edge of

fast-flowing mountain streams but is also present in the vicinity of still waters. A. khasiense has also been found in

a ricefield located at the edge of a disturbed primary forest or seen active along a man-made pool surrounded by

agricultural fields inhabited by newts of the species Tylototriton verrucosus Anderson, 1871 and various frogs.

This active, terrestrial and semi-aquatic species is both diurnal and nocturnal. It lives mainly among leaves of the

forest litter and in the vegetation surrounding of pieces of water. This snake feeds mainly on small frogs, tadpoles

and, perhaps, insects. Wall (1926) recorded one or two frogs from the stomach of 12 specimens. This species is

oviparous; Wall (1926) removed from 1 to 4 eggs from his series of specimens; Whitaker and Captain (2004)

recorded new born specimens in June and October but nothing else is known on its reproductive habits.



FIGURE 3. One of the syntypes of Tropidonotus khasiensis, BMNH 1946.1.12.81, Khasi Hills, India. Photos Gernot Vogel.



FIGURE 4. Comparison of the pattern of the upper labials of three species of the Amphiesma khasiense group. A. A. boulengeri, BMNH 1969.1710, Kampot Province, Cambodia. B. A. boulengeri, FMNH 180153, Nakhon Ratchasima Province, Thailand. C. A. modestum, BMNH 1925.9.17.2, Kachin Hills, Myanmar. D. A. khasiense, BMNH 1946.1.12.82, syntype, State of Meghalaya, India. E. A. khasiense, BMNH 1946.1.13.63, holotype of Natrix gilhodesi, Kachin State, Myanmar. F. A. khasiense, BMNH 1925.4.2.13, Kachin State, Myanmar. Photos Gernot Vogel.

Amphiesma boulengeri (Gressitt, 1937)

(Fig. 4A–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—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, 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 63II5098, 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 0002 S,

Tam Dao Hill Station. Quang Nam Province. AMNH 148552, Tra My; IEBR 2977, ROM 03848, 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 07484, 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.



FIGURE 5. Comparison of the pattern in life of four species of the Amphiesma khasiense group. Top left. Amphiesma khasiense (Boulenger, 1890), Umphang, Tak Province, Thailand. Photo Sjon Hauser. Top right. Amphiesma khasiense (Boulenger, 1890), Nokrek Biosphere Reserve, State of Meghalaya, India. Photo Ashok Kumar Mallik. Centre left. Amphiesma boulengeri (Gressitt, 1937), Hainan Island, China. Photo Kevin Messenger. Centre right. Amphiesma boulengeri (Gressitt, 1937), Phnom Aural Province, Cambodia. Photo Lee Grismer. Bottom. Amphiesma boulengeri (Gressitt, 1937), Kon Tum Province, Vietnam. Note the dorsal spots and separate blotches on supralabials, closer to the pattern of Amphiesma khasiense rather than to a typical A. boulengeri. Photo Truong Quang Nguyen.



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.

TABLE 3. Main variation in selected meristic characters in Amphiesma boulengeri.

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

Sex / Cluster Ventrals Subcaudals DSR Red. 1 DSR Red. 2

Males

Cluster I 140–152x = 146.0s = 3.1

93–109x = 102.5s = 3.9

85–97x = 87.3s = 3.8

85–99x = 89.60s = 3.8

Cluster II 136–159x = 145.3s = 6.1

84–101x = 95.7s = 4.3

81–91x = 85.5s = 4.1

83–97x = 89.0s = 4.8

Cluster III 155–165x = 160.7s = 3.1

108–119x = 113.5s = 4.5

86–100x = 95.7s = 4.6

87–103x = 97.7s = 5.2

Females

Cluster I 139–151x = 144.0s = 3.4

90–100x = 95.6s = 3.6

82–89x = 84.0s = 3.4

83–90x = 86.0s = 2.7

Cluster II 136–153x = 143.8s = 5.6

83–101x = 89.3s = 4.7

82–97x = 87.2s = 5.5

86–103x = 90.7s = 4.7

Cluster III 152–166x = 157.9s = 5.7

99–115x = 108.5s = 5.7

92–100x = 95.6s = 5.7

98–103x = 99.1s = 5.7



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 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.

Amphiesma inas (Laidlaw, 1901)

(Fig. 6)

Tropidonotus inas Laidlaw, 1901: 576, Pl. 35: Fig. 3 & 4. – Type locality. “Gunong Inas”, now Gunung Inas [Mt. Inas], near Lawin, State of Perak, West Malaysia, Federation of Malaysia. – Holotype. A juvenile, currently not traced (not in BMNH or ZRC collections); collected by Messrs. Laidlaw and Yapp, Skeat Expedition, 1899–1900.

Tropidonotus inas.—Boulenger 1912: 123 & 125—Smith 1916: 159—Werner 1929: 25Natrix inas.—Smith 1930: 43; 1943: 291 (specimens from “Pen. Siam and Malay Peninsula”)—Smedley 1932: 115—Pope

1935: 90—Bourret 1936a: 103, 113 & 132.—Tweedie 1953: 70, 71: Fig. 17a-b, 65, 123; 1957: 72, 73: Fig. 18a-b, 67, 127.Amphiesma inas.—Malnate 1960: 50, 52 & 57—Taylor 1965: 821, 829 & 830: Fig. 54—Tweedie 1983: 86, 93 & 94: Fig. 22a-

b, 156—Welch 1988: 30—Cox 1991: 235, 236: Fig. 21—Manthey & Grossmann 1997: 304 & 313—Cox et al. 1998: 44—Chan-ard et al. 1999: 30 & 153 (center and bottom)—Chanhome et al. 2001: 58.—Inger & Voris 2001: 887—Iskandar & Colijn 2001: 96 (in part)—David & Das 2003: 416, 417 & 418—Leong & Lim 2003: 132—Nabhitabhata et al. 2004: 124 (in part: mentions of Khao Luang, Nakhon Si Thammarat Province, and of Prachuap Khiri Khan)—Pauwels et al. 2009: 77—Das 2010: 150: Pl. 67, 332.

Material examined (10 specimens). Thailand. Nakhon Si Thammarat Province. BMNH 1916.3.27.31, Khao

Wang Hip (upper camp).—Federation of Malaysia. West Malaysia. State of Pahang. MNHN 1999.9092, Cameron

Highlands.—ZRC 2.4055, Cameron Highlands, 4,500′; ZRC 2.4056–4058′, Cameron Highlands, 4–5,000′.—ZRC

2.4059, Fraser’s Hill.—ZRC 2.5920, Gunung Brinchang.—BMNH 1938.8.7.13–14, Bukit Fraser (or Fraser’s

Hills).

Taxonomic comments. Amphiesma inas was described from a juvenile specimen which cannot be currently

traced. In contrast to the statement of Iskandar & Colijn (2001), this type is not in the collections of the Natural

History Museum of London. As defined here, A. inas is endemic to Peninsular Thailand and West Malaysia. This

species is monotypic.

Diagnosis. A species of the genus Amphiesma characterized by the combination of (1) a moderately elongate

body; (2) nostrils directed laterally; (3) 19–21 maxillary teeth, gradually enlarged, the last 2 teeth distinctly



enlarged; (4) 19 dorsal scale rows at midbody, strongly keeled on upper rows, moderately but distinctly keeled on

1st scale row; (5) dorsal pattern made of a series of yellow or ochre (cream in preservative) dorsolateral spots on a

faint dorsolateral stripe (or absent), the first two or three enlarged and connected by a yellow stripe, other also

enlarged on the anterior part of the body; (6) each posterior supralabial with a large, elongate, straight or oblique,

cream blotch, or scale largely cream; (7) pattern of the neck made of a conspicuous, pale stripe extending from the

corner of the mouth to the first enlarged dorsolateral spot; (8) venter ivory or cream, with the outer quarter or third

of each ventral dark reddish-brown, in contact with the dorsal colour; (9) eye large, 1.9–2.0 times the distance

between the lower margins of eye and of lip; (10) 141–151 VEN, 93–109 SC; (11) internasals abruptly truncated

anteriorly, (12) 9 supralabials; and (13) 1 anterior temporal.

FIGURE 6. Living specimen of Amphiesma inas (Laidlaw, 1901), San Yen, Khao Nan National Park, Nakhon Si Thammarat Province, Thailand. Photo Michael Cota.

Amphiesma inas is easily distinguished from A. boulengeri by (1) 2nd to 9th DSR more keeled in A. inas, (2)

scales of 1st DSR keeled (vs. smooth in A. boulengeri), and (3) a distinct pattern on posterior supralabials. In

contrast, A. inas and A. khasiense are very similar. The most obvious characters for separating these two species are

the dorsolateral spots, yellow and conspicuous in A. inas, the first two or three distinctly enlarged and connected by

a wavy stripe, small and rather reddish-brown in A. khasiense, and the stripe on the neck, broad in A. inas and

connected to the first enlarged dorsal spot, short, narrow or often broken into spots in A. khasiense. Characters of

use for separating these taxa are given below in the Discussion.

Variation (based Smedley 1932, Taylor 1965, Tweedy 1983 and on 8 examined specimens). Body

cylindrical, elongate, rather slender in males, slightly more robust in old females, slightly laterally compressed;

head rather elongate, oval, depressed in front of the eye, distinct from neck; snout rather short, blunt, amounting for

25.0–27.5 % of HL or 1.30–1.55 times as long as diameter of eye; nostrils large, crescentic, directed laterally and

piercing in the middle of the nasal; eye large, 1.90–2.05 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 known is 634 mm (SVL 428 mm; TaL 206 mm; specimen ZRC 2.5920; male). The



SVL of the longest known female is equal to 549 mm, which would give a TL of about 670 mm. Ratio TaL / TL:

0.321–0.342, without sexual dimorphism.

Dentition. Maxillary teeth 19–21, gradually enlarged, the last 2 teeth distinctly enlarged.

Body scalation. DSR: 19–19–17 scale rows; dorsal scales rhomboedric, normal or notched at their posterior

extremity, distinctly keeled, more strongly in the posterior half of the body on all rows; scales of 1st DSR more or

less strongly keeled, never smooth.

Scale row reductions: first reduction (19→17) at VEN 85 (at left) and second reduction (17→15) at VEN 87

(at left) in a single specimen.

VEN: 141–151 (plus 1 preventral in all examined specimens); SC: 93–109, all paired, without sexual

dimorphism; ratio VEN / SC: 1.35–1.50; anal plate divided.

Position of the reduction to 6 scale rows around the tail: 11th–21st SC, with a sexual dimorphism (see

dimorphism). Ratio length of the portion of tail with 4 dorsal scale rows / length of the portion of tail with 6 dorsal

scale rows: 1.20–1.50.

Head scalation. Rostral trapezoidal wider than high, barely visible from above; nasals subrectangular,

elongate, vertically divided on their lower half, with the posterior part larger and higher than anterior one;

internasals subtriangular, rather small, in broad contact, about 1.1–1.2 times as long as wide, abruptly truncated

anteriorly with anterior margin about 0.45–0.60 times the width of the posterior margin; 2 wide, subrectangular

prefrontals, 1.3–1.5 times as long as internasals; frontal hexagonal, shield-like with apex directed posteriorly, rather

small, 1.35–1.55 longer than wide, 1.8–1.9 times as long as prefrontal; 1 supraocular on each side, longer than

wide, about as wide as internasals; two large parietals, 1.2–1.4 times longer than frontal; 1 small, rectangular loreal

scale, 0.8–1.3 times as high as long, in broad contact with the nasal; 9 SL, 3rd–9th longer than high; 1st and 2nd SL

small and short, in contact with nasal; 2nd and 3rd SL in contact with the loreal; 4th–6th SL entering orbit; 7th and 8th

SL largest; 1 relatively large or 2 small preoculars on each side; 2 or 3 small postoculars; 1 anterior temporal,

narrow and elongate, with complete temporal formulas as 1 + 2 + 2; 10 infralabials (9 in 1 / 16 occurrences), first

pair in contact each with other, 1st–4th IL in contact with anterior chin shields, 6th IL largest.

Coloration and pattern in alcohol. The upper surface of body is dark reddish-olive brown, dark greyish-brown,

dark reddish-brown or blackish-brown, variegated with darker pigmentation producing 2 or 3 rows of faint,

irregular blotches on the upper scale rows and on the sides; a well defined pale yellow or pale grey dorsolateral

stripe, wavy or broken by two or three large yellow blotches, extends on 5th and 6th DSR from the neck and

foremost part of the body, turning into a pale, irregular, creamish-brown or rusty brown stripe extending on upper

half of 5th and the whole of 6th dorsal scale rows, sometimes also on lower edge of 7th DSR, progressively fainter,

usually not visible after midbody; on this dorsolateral stripe or on the background colour, a series of cream or

yellow (coral, orange or red in life) spots, about 1 scale long and at 2–4 scales from each other, rather large and

conspicuous on the anterior quarter or third or body, becoming smaller posteriorly; some very narrow cream dashes

between dorsolateral spots on the anterior third of the body; a short, narrow ventrolateral stripe extending on the

lower edge of scales of 1st DSR up to about the level of 12th to 30th VEN. The tail is as the body, with minute

dorsolateral cream or yellow (coral, orange or red in life) spots, vanishing progressively and disappearing

completely at mid-length of the tail; posterior half of tail uniformly dark, variegated with irregular and faint darker

pigmentation.

The head is dark greyish-brown or reddish brown, paler than the background colour of the body, with irregular

dark brown vermiculations; upper head surface with irregular paler areas, speckled with minute blackish-brown

dots; usually a small, yellow spot on the each parietal; a short cream streak just behind parietals present or absent;

anterior six supralabials dark with an irregular cream spot, larger on 5th and 6th supralabials reducing the dark colour

to the posterior edge of the scale; 7th, 8th and 9th supralabials blackish-brown, darker than upper head surface, with

either a large blotch in their middle, pure white or cream (same in life), slightly oblique, horizontally elongate, or

with a triangular blotch covering the lower part and middle of the supralabial, in both cases not in contact with each

other, contrasting sharply with the dark colour of the head; blotch more irregular in 9th SL; no postocular streak; on

each side of the neck, either a conspicuous series of rounded or elongate cream, yellow or pale grey blotches, or a

rather wide but continuous, oblique, cream, yellow or pale grey stripe extends obliquely from the corner of the

mouth to the anterior part of the dorsolateral stripe or to the first dorsolateral spot. The throat and chin are cream or

ivory; infralabials and mental usually heavily spotted with small, irregular dark reddish-brown spots; usually

minute dots on the preventral area.



The venter is uniformly ivory or cream (yellow in life), with about 1/4 to 1/3 of the outer part of each scale

very dark brown, in contact without discontinuity with the background colour of the body except for a short

distance on the anterior body; sometimes a minute, elongate cream spot on the outer, posterior edge of each ventral

just at the level of the base of ventral tips, producing a narrow, discontinuous ventral stripe separating the dark tip

from the dark ventral blotch. The tail is ivory or cream below, with outer edges of subcaudals very dark brown;

posteriorly, the tail becomes darker.

Hemipenis. In situ, it is short and stout as in other species of the group, 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. The organ is entirely covered with short but dense spines, at the exception of the

base which is covered with large calyces; lips of the sulcus also densely covered with short, thick spines; 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. In our limited sample, we could find a sexual difference only in the position (counted in

number of subcaudals) of the reduction from 6 to 4 scale rows around the tail of subcaudals, with males: 19th–21st

SC vs. females: 11th–15st SC.

Distribution.—According to the definition of Amphiesma inas given here and authors cited above: Thailand.

Prachuap Khiri Kahn Province. Huay Tam Phra (after Taylor 1965; specimen not seen). Nakhon Si Thammarat

Province. Khao Wang Hip; San Yen, Khao Nan National Park, 08° 46' N 099° 31' E, ca. 1,300 m a.s.l.; Fig. 6).—

Federation of Malaysia. West Malaysia. State of Perak. Gunung Inas. State of Pahang. Cameron Highlands; Bukit

Fraser (Fraser’s Hills); Genting Highlands; Gunung Brinchang. This species should be searched in other mountain

ranges of West Malaysia.

In the literature, Amphiesma inas has also been cited from the island of Sumatra in Indonesia (Mumpuni 2001).

We have examined the specimen from Sumatra (MZB 2186). It proved to be the second known specimen of

Amphiesma kerinciense David & Das, 2003 (see below). Lastly, Das (2010) mentioned Amphiesma inas from

several Thai provinces as far north as Loei Province. All specimens of this group recorded from Thailand north of

the Isthus of Kra and seen by us are referable to Amphiesma khasiense or A. boulengeri.

Biology. A. inas has been recorded only from forested areas at high elevations between 1,000 and 1,500 m a.s.l.

According to Das (2010), it is nocturnal and terrestrial, active on the forest litter and among buttresses. Little else is

known about the biology of this seemingly rare species.

Amphiesma kerinciense David & Das, 2003

(Fig. 7)

Amphiesma inas (nec Tropidonotus inas Laidlaw, 1901, a valid species).—Mumpuni 2001: 38—David & Das 2003: 417 (citation of the specimen described by Mumpuni 2001).

Amphiesma kerinciense David & Das, 2003: 414, 415: Fig. 1–4.—Type locality. Sungai Jalnei Dalam, at base of Gunung Tugu (or Tujuh) [= Mt. Tugu or Tujuh] (01°42'59.0"S-101°21'43.1"E), Gunung Kerinci [Mt. Kerinci], Sumatera Barat Province, Sumatra Island, Indonesia.—Holotype. ZRC 2.3521, adult female; collected by Darren Yeo and Heok Hui Tan, 12 June 1996.

Amphiesma kerinciense.—Das 2010: 332.

Material examined (2 specimens). Indonesia. Sumatra. Lampung Province. MZB 2186, Kubu Perahu, Bukit

Barisan Selatan National Park, near Lake Ranau. Sumatera Barat Province. ZRC 2.3521 (holotype), Sungai Jalnei

Dalam, at base of Gunung Tugu (or Tujuh) (01°42'59.0"S-101°21'43.1"E), Gunung Kerinci.

Taxonomic comments. Amphiesma kerinciense was described from a single, female specimen. Mumpuni

(2001) briefly described a specimen of Amphiesma from the southern Barisan Range (MZB 2186) which she

identified as A. inas. After examination, this specimen proved to be referable to Amphiesma kerinciense. We here

slightly expand the description of this species on the basis of the second known specimen and the first known male.

Amphiesma kerinciense seems to be allied to the Indo-Malayan Amphiesma inas but its affinities may also lie

with Amphiesma sanguineum (Smedley, 1932).

Diagnosis. A species of the genus Amphiesma characterized by the combination of (1) a stout body; (2) nostrils

directed laterally; (3) 20–21 maxillary teeth, gradually enlarged, the last 2 teeth distinctly enlarged; (4) 19 dorsal



scale rows at midbody, strongly keeled on upper rows, many of which are distinctly notched posteriorly; scales of

first dorsal row enlarged, feebly keeled; (5) dorsal pattern made of a broad, darker vertebral band, with or without

irregularly placed dark scales producing faint and irregular, discontinuous crossbars, edged on each side by a faint

pale ochre brown dorsolateral stripe, widening to produce three or four irregular blotches on the neck; (6) posterior

supralabial slightly and irregularly paler, without definite blotches; (7) neck uniform or with a narrow pale streak;

(8) venter pale with a row of well defined dark brown blotches at 3/4 of their width; (9) eye large, about 2.0 times

the distance between the lower margins of eye and of lip; (10) 138–140 VEN; (11) internasals broadly truncated

anteriorly, (12) 8 or 9 supralabials; and (13) 1 anterior temporal.

FIGURE 7. Head portrait of Amphiesma kerinciense David & Das, 2003, ZRC 2.3521, holotype, adult female. Photo Patrick David.

Variation (based on 2 specimens). Body stout, cylindrical, elongate; head rather elongate, oval, depressed in

front of the eye, barely distinct from neck; snout long, blunt when viewed from above, rectangular in lateral view,

amounting for about 26.0 % of HL or 1.6 times as long as diameter of eye in two specimens; nostrils large,

crescentic, directed laterally and piercing in the middle of the nasal; eye large, about 2.0 times the distance between

the lower margin of eye and the lower edge of lip, with a round pupil; tail long, cylindrical and tapering.

The maximal total length known is 516 mm (SVL 358 mm; TaL 158 mm; specimen ZRC 2.3521; female,

holotype). The SVL of the second available specimen, a juvenile male with an incomplete tail, is only 277 mm.

Ratio TaL / TL: 0.308 in the female specimen.

Dentition. Maxillary teeth 20–21, gradually enlarged, the last 2 teeth distinctly enlarged.

Body scalation. DSR: 19–19–17 scale rows; dorsal scales rhomboedric, usually distinctly notched at their

posterior extremity, strongly keeled with a narrow, sharp keel; dorsal scales, more strongly in the posterior half of

the body on all rows; scales of 1st DSR feebly keeled or nearly smooth.

VEN: 138–140 (plus 0 or 2 preventrals); SC: 89 in the adult female, all paired; ratio VEN / SC: 1.56; anal plate

divided.

Position of the reduction to 6 scale rows around the tail at the level of 8th SC. Ratio length of the portion of tail

with 4 dorsal scale rows / length of the portion of tail with 6 dorsal scale rows: 1.45.



Head scalation. Rostral trapezoidal wider than high, visible from above; nasals subrectangular, elongate,

vertically divided on their lower half, with the posterior part larger and higher than anterior one; internasals

subtriangular, rather small, in broad contact, about 1.0 to 1.2 times as long as wide, abruptly truncated anteriorly

with anterior margin about 0.5-0.6 times the width of the posterior margin; 2 wide, subrectangular prefrontals, 1.6–

1.8 times as long as internasals; frontal hexagonal, shield-like with apex directed posteriorly, rather small, 1.4 times

longer than wide, 1.9–2.0 times as long as prefrontal; 1 supraocular on each side, longer than wide, about 0.3 times

as wide as frontal, narrower than internasals; two large parietals, 1.2–1.4 times longer than frontal, in contact for a

length 1.1 times as great as the frontal length; 1 small, rectangular loreal scale, 0.8 times as high as long, in broad

contact with the nasal; 8 or 9 SL, 2nd–8th or 9th longer than high; 1st and 2nd SL small and short, in contact with nasal;

2nd and 3rd SL in contact with the loreal; 4th–5th or 4th–6th SL entering orbit; 6th and 7th SL or 7th and 8th SL largest; 1

relatively large or 2 small preoculars on each side; 3 postoculars, the upper one largest; 0 or 1 subocular; 1 anterior

temporal, narrow and elongate, with complete temporal formulas as 1 + 1 + 2 or 1 + 2 + 2; 10 or 11 infralabials,

first pair in contact each with other, 1st–4th or 1st–5th IL in contact with anterior chin shields, 6th IL largest; mental

scale small; posterior chin shields about 1.2 times longer than anterior ones.

Coloration and pattern in alcohol. Background colour of sides, up to 4th dorsal scale row row, dark reddish

grey-brown, due to an intricate speckling of dark reddish-brown pigments on a pale greyish-ochre background,

with 1st–2nd lower scale rows rather purple greyish-brown, slightly iridescent, with ventroposterior extremities of

scales of the 1st row dark brown; widely scattered small blackish-brown spots on 2nd, 3rd and 4th dorsal rows; these

dark brown spots produce faint and irregular, discontinuous crossbars, more visible on the posterior part of the

body. A broad blackish-brown vertebral band on 5th–9th scale rows and the vertebral row, with indistinct irregular

black crossbars. A dorsolateral stripe extending from the neck to the tail on top of 5th and the whole of 6th scale row,

well defined and greyish-pink on the neck and foremost part of the body (up to 6th VEN), becoming discontinuous

and broken into 4 or 5 irregular and diffuse blotches more or less connected, then changing into a diffuse, ill

defined pale ochre-brown stripe. The tail is blackish-brown above, greyish-ochre on the upper part of the side in the

extension of the dorsolateral band of the body, with a blackish-brown band on its lower side and extremities of

subcaudals, divided into two equal parts by a narrow greyish-purple line in the extension of the colour present on 1st

and 2nd scale rows of the body.

Head dark reddish grey-brown above with ochre vermiculations, turning into blackish-brown on its posterior

part; a fine speckling with purple greyish-brown on temporals; two small faint yellowish-white spots on the

parietals and a very faint whitish-brown sagittal line just behind the parietals. Anterior supralabials are ochre

brown, with brown speckling and edged with dark brown on their posterior part. Posterior supralabials light ivory

cream; posterior part of 6th SL blackish-brown; 7th SL with a curved dark brown marking in the posterior part of its

centre and with its posterior upper corner dark brown; 8th SL divided into a lower anterior part ivory cream and an

upper posterior part dark reddish grey-brown, these two parts being separated by an oblique blackish-brown streak

connecting the upper tip of the 7th SL to the lip edge at the junction of 8th and 9th SL; 9th SL dark reddish grey-

brown, with its lower part blackish-brown and its upper part ochre brown, narrowly bordered with blackish-brown

above. An ill defined and narrow postocular streak, pale ochre brown narrowly edged below with blackish-brown,

extending from the upper preocular to 9th SL through the anterior temporal and the upper tip of 8th SL. Behind the

9th SL, a narrow, irregular pale postocular streak, bent upwards; on the side of the neck, a very thin, greyish-pink

stripe, connected both to the postocular stripe and the dorsolateral stripe. Throat, chin and ventral part of the neck

ivory cream, uniform in their middle, with some dark brown speckling on the outer parts of the throat and lower

side of the neck; posterior margins of infralabials dark brown.

Venter uniformly ivory cream, with about 1/4th of the outer part of each scale purple greyish-brown as dorsal

rows 1–2, becoming blackish-brown on the scale outer tip; a subrectangular, irregular blackish-brown spot between

the purple greyish-brown and ivory parts; these spots become progressively larger backwards and are connected to

the purple greyish-brown colour of the flank from about the 20th VEN onwards; before this point, the blackish-

brown ventral spots are separated from the purple greyish-brown colour of the lower flank by a narrow ivory cream

line, producing a thin and conspicuous but discontinuous series of dark brown spots at the bottom of the flanks; the

ivory line is progressively replaced by the purple greyish-brown hue. Below, the tail is as the venter, bordered on

each side by a blackish-brown stripe described above, with a middle line made of irregular and faint dark brown

speckling.



Hemipenis. Unknown.

Sexual dimorphism.Unknown.

Distribution. Indonesia. Sumatra. Endemic to this island. Province of Sumatera Barat. Massif of Mt. Kerinci.

Lampung Province. Bukit Barisan Selatan National Park, near Lake Ranau.

Biology. The holotype of A. kerinciense was collected in a shallow hill stream, less than 30 cm deep, fast

flowing in parts through an open, grassy area. The elevation was unfortunately not recorded. The clear water

flowed over a substratum of stone and gravel. The snake was collected while it was feeding on tadpoles, possibly of

the genus Huia, which it later regurgitated. Nothing else is known on the biology of this rare species.

Discussion

A summary of the confusions. The overall similar external morphology of the three mainland species, A.

khasiense, A. boulengeri and A. inas, has led to an intricate tangle of names in the literature. In our opinion, this

confusion arose mostly because the three main reference books dealing with the region encompassing southern

China and Indochina, i.e., Pope (1935), Bourret (1936a–b), and Smith (1943) were not aware of the existence of

Amphiesma boulengeri (Gressitt, 1937). A second reason comes from the comparatively short descriptions of

Tropidonotus khasiense Boulenger, 1890 and Tropidonotus modestus Günther, 1875, which were not sufficient to

distinguish between these very similar species.

Pope (1935: 90) noted that specimens referred to Tropidonotus modestus by Mell (1922: 116) then to “Natrix

khasiensis>gilhodesi” [sic, the meaning of the „> “ is not clear to us] (Mell 1931: 203) were distinct from the Indo-

Himalayan A. khasiense and A. modestum. Furthermore, Pope noted that specimens from Langbian Plateau,

southern Vietnam, cited by Smith (1921) as Tropidonotus johannis (see below), probably belonged to the same

species than Mell’s specimens. For some reasons, Pope could not secure any specimen of A. boulengeri during his

own investigation.

Bourret (1936b) rightly recognized the existence of several taxa of this group in Vietnam but, lacking

comparative material of genuine A. khasiense from the Indo-Himalayan Region, he confused specimens now

referred to A. boulengeri with the morphologically closest species, A. khasiense. This misidentification has been

repeated from Smith (1943) up to recent times, in publications, for example in Nguyen et al. (2009) and Das

(2010). The eastern limit of distribution of A. khasiense is in Phongsaly Province, northern Laos. Although A.

khasiense might well occur in mountains of north-western Vietnam, we have yet to see a genuine A. khasiense from

this latter country. All Vietnamese specimens previously identified as A. khasiense examined by us are typical A.

boulengeri. Furthermore, the mentions and descriptions given by Bourret (1936a-b) under the combinations A.

modesta or A. modesta modesta, are either referrable to A. modestum, a species not present in Vietnam, or,

seemingly, to A. boulengeri for a part, and even to a third species under description, indeed related to A. modestum

and present in northern Vietnam. This yet undescribed species is described and even depicted in Bourret (1936b:

72–73; 74: Fig. 30). It is morphologically close to the genuine A. modestum but differs by characters of which

Bourret could not be aware due to a lack of comparative material. Lastly, Bourret (1936a–b), following Smith

(1921), referred specimens from the Plateau of Langbian, in Dalat Province, to as Tropidonotus johannis

Boulenger, 1908 or Natrix modesta johannis. This Chinese species, now Amphiesma johannis, belongs to the

Amphiesma khasiense species-group but can be identified by characters given in the key below.

Smith (1943) correctly described Amphiesma khasiense but, following Bourret (1934a, 1935a–c), referred to

this species specimens from Tam Dao, in northern Vietnam. Although Smith’s book was published in late 1943, it

was completed in 1938 according to author’s preface. Either Smith could not see the description of A. boulengeri,

or probably, neglected it because it was then treated as a purely Chinese species. Furthermore Smith (1943)

adopted quite a large and rather crude definition of A. modestum in including under this combination no less than

three valid species, i.e. A. modestum, A. deschauenseei (Taylor, 1934), and A. boulengeri for populations cited on p.

291 from “Upper Laos (Chieng Khoung [sic])”, “Cambodia (Kamchay Mts.)” and “S. Annam (Langbian Plateau)”.

In contrast to Smith (1921), Smith (1943) referred specimens from the Langian Plateau to A. modestum. Campden-

Main (1970) followed this position and also mentioned A. modestum from South Vietnam. Lastly, following Smith

(1916), Smith (1943) correctly referred to Natrix inas, now A. inas, specimens of populations from “Pen. Siam and

Malay Peninsula”, i.e. to Peninsular Thailand and West Malaysia. We examined the specimen from “Peninsular



Siam”, seen by Smith (1916), i.e., from Nakhon Sri Thammarat Province (BMNH 1916.3.27.31). It is a typical

Amphiesma inas. However, Smith suggested that Tropidonotus inas Laidlaw, 1901 may be only a “racial form” of

A. modestum. Both species do not belong to the same group and are easily distinguished by characters given below.

As a summary, in the literature the four species discussed here have been identified or cited as follows:

- Amphiesma khasiense (Boulenger, 1890), as defined here, has always been correctly identified at the exception

of Chan-ard et al. (1999: 153; top, specimen from Phu Luang, Loei Province, Thailand, misidentified as A. inas)

and Nabhitabhata et al. (2004: 124; also record of Phu Luang, Loei Province), and Cox et al. (1998) and Das

(2002), in both case misidentified as A. modestum, and cited under the various combinations listed in the

chresonymy above or as its junior synonym Natrix gilhodesi Wall, 1925. Without justification, Mahendra (1984)

synonymized A. khasiense with A. modestum.

The specimen from Phu Luang, Loei Province, depicted by Chan-ard et al. (1999: 153) was erroneously

identified by these authors as A. inas. It is indeed an Amphiesma khasiense, in spite of its rather elongate blotches

on the posterior supralabials. The lack of dorsolateral stripes excludes its identification as A. boulengeri.

Nabhitabhata (1987) listed, without comments nor voucher material, Amphiesma modestum from Doi Suthep-Pui

National Park, Chiang Mai Province, northern Thailand. That record might refer to Amphiesma khasiense, or, more

likely, to A. deschauenseei, a valid species related to A. modestum.

- Amphiesma boulengeri (Gressitt, 1937), as defined here, has largely been correctly identified and cited in

publications dealing with China from Gressitt (1937) onwards. Previously, it had been cited as Tropidonotus

modestus by Werner (1904), Mell (1922) and Vogt (1922) or Natrix khasiensis gilhodesi by Gée (1930). In contrast,

misidentifications and erroneous combinations are the rules for populations of the Indochinese Peninsula in most

publications from Smith (1943) onwards. In chronological order, this species has variously been cited (see above

for the authors) as:

Vietnam: 1) Amphiesma modestum, through various combinations.—2) Amphiesma johannis, through various

combinations, following Smith (1921), only for the population of Langbian Plateau.—3) Amphiesma khasiense,

widely cited under the combinations Natrix khasiensis and Amphiesma khasiensis following the erroneous

identifications of Mell (1929) and Bourret (1934a, 1936b), upon which are seemingly based the numerous,

erroneous records and citations of Amphiesma khasiense in Vietnam until Chanhome et al. (2001) and David et al.

(2007).

Laos: 1) Amphiesma modestum, as Natrix modesta (Angel 1929; Smith 1943).—2) Amphiesma khasiense, as

Natrix khasiensis (Deuve 1970).

Cambodia: 1) Amphiesma modestum, as Natrix modesta by Smith (1943).—2) Amphiesma parallelum, as

Natrix parallela parallela, by Bourret (1934b, 1936b). No locality is cited and we could not examine these

specimens but, on the basis of the detailed description provided by Bourret (1934b), these specimens are referrable

to Amphiesma boulengeri. Amphiesma parallelum is an Indo-Himalayan species unknown from Cambodia and

from the Indochinese Peninsula as a whole. The related species Amphiesma bitaeniatum (Wall, 1925) has been

recorded from mountains of northern Vietnam but is unknown in Cambodia (David et al. 2005).—3) Amphiesma

cf. khasiense by Grismer et al. (2007).

Thailand: 1) Amphiesma modestum, as Natrix modesta by Smith (1943).—2) Amphiesma inas, by

Nabhitabhata et al. (2004; record of Khao Yai, Nakhon Ratchasima Province).

- Amphiesma inas (Laidlaw, 1901), as defined here, has always been correctly identified as far as specimens

from South Thailand West Malaysia are concerned. Records from northern Thailand refer to A. khasiense.

- Amphiesma kerinciense David & Das, 2003, had been misidentified as A. inas by Mumpuni (2002).

Morphological comparisons between the four species. These four species are morphologically so close that

they can be positively identified only in using a combination of characters. The locality of collect of the specimens

to be identified will greatly help but cannot be considered as a character in itself to differentiate nearly sympatric or

sympatric taxa such as A. khasiense and A. boulengeri (especially in north Laos and Yunnan) or A. boulengeri and

A. inas (in Peninsular Thailand). We provide in Table 4 a series of characters, the combination of which should

allow their identification.



TABLE 4. Main characters allowing the identification of Amphiesma khasiense, A. boulengeri, A. inas and A.

kerinciense. Abbreviations (from left to right). MT: number of maxillary teeth. K1SR: keeling of scales of the 1st dorsal

scale rows. Pattern of posterior supralabials. IRB: isolated rounded blotches; IOB: isolated ovale or subrectangular

blotches; IHS: isolated horizontal streaks; CHS: contiguous horizontal streaks; CST: continuous stripe. Pattern of neck.

UNI: uniform, no pattern; FRD: feature (stripe, series of blotches or streak) reaching the dorsum; IBL: isolated blotches;

BLC: blotches connected; NST: narrow stripe; BST: broad stripe. Dorsal pattern. DST: dorsal stripe present (at least on

anterior part of the body); FST: dorsal stripe faint; BST: dorsolateral stripe broad and conspicuous; DSP: dorsal spots

present; SSP: dorsolateral spots small; LSP: dorsolateral spots large, conspicuous. VEN: ventral scales. SC: subcaudal

scales.

TABLE 4. (Continued)

Before discussing the differences between these species, we here provide an updated diagnosis of Amphiesma

modestum (Günther, 1875) (Fig. 23–25). This Indo-himalayan species has often been confused with members of

the A. khasiense group and is also currently misidentified in the literature. On the basis of our data, we suggest: (1)

19 dorsal scale rows at midbody, (2) nostrils directed dorsolaterally, not as lateral as in the A. khasiense group, (3)

dorsal scales feebly keeled, scales of 1st row entirely smooth, (4) upper scales at the base of the tail not strongly

keeled, (5) tail long, ratio TaL/TL 0.288–0.326, (6) 104–117 SC, (7) 9 supralabials, (8) supralabials either nearly

entirely pale, speckled with dark brown or more or less dark brown or blackish-brown, with vague paler areas, but

never with defined blotches, (9) no postocular, (10) a wide, horizontale pale stripe on the neck, (11) a broad but

more or less visible dorsolateral stripe on each side, and (12) venter pale brown or ochre in its middle, more or less

extensively dark on the sides. Furthermore, A. modestum is known only from north-eastern India, northern

Myanmar and extreme south-western China (western Yunnan Province). Specimens with a black venter or broad

dorsolateral blotches should be referred to other species, not treated here; a revision of this group is in progress.

A. kerinciense, with its subdued dorsal and labial patterns, is the easiest species to recognize, although a casual

examination may lead to a misidentification with A. modestum. However, A. kerinciense shares most scalation

characters with Amphiesma inas (Laidlaw, 1901) and A. sanguineum (Smedley, 1932). This latter species is

endemic to mountains of West Malaysia. Amphiesma kerinciense and Amphiesma sanguineum share a similar

dorsal pattern made of a broad, dark vertebral band extending on rows 7-10, ornated with black markings, and most

characters of the body and head scalation. Main differences are (1) a higher number of maxillary teeth in A.

kerinciense, 18 or 19 +2 enlarged teeth without diastema, vs. 15+2 with a small diastema in A. sanguineum, (2)

dorsal scales notched posteriorly in A. kerinciense, normal in A. sanguineum, (3) a different dorsal colour, greyish-

brown or brown in A. kerinciense, reddish-brown in A. sanguineum; (4) paler supralabials in A. kerinciense, ivory

vs. ochre yellow in A. sanguineum; (5) a different pattern on posterior supralabials, with a broader oblique dark

brown streak in A. sanguineum; and (6) centre of throat and chin more or less speckled with dark brown spots in A.

sanguineum.

Diagnostic differences between Amphiesma khasiense, A. boulengeri and A. inas were presented in Chanhome

et al. (2001) and David et al. (2007) but are here precised or slightly corrected. As far as A. khasiense, A.

boulengeri and A. inas are concerned, the most useful external characters are the pattern of the posterior

Species MT K1SR Pattern of posterior supralabials Pattern of neck

IRB IOB IHS CHS CST UNI FRD IBL BLC NST BST

A. khasiense 21–26 + + + – / + – – – – + – – (+) –

A. boulengeri 25–28 – – – + + + – + – + + +

A. inas 19–21 + + + – – – – + + + + +

A. kerinciense 20–21 + – – + – – – + – – + –

Species Dorsal pattern VEN SC

DST FST BST DSP SSP LSP

A. khasiense – / + + – + + – 141–155 87–111

A. boulengeri + – + + + – 136–166 83–115

A. inas – (+) + – + + + 141–151 93–109

A. kerinciense + + + – – – 138–140 89



supralabials, the pattern on the neck, the presence or not of a conspicuous dorsolateral stripe and the size of the

dorsal dots, although none of these characters taken alone are sufficient. A. khasiense and A. inas have (1) more or

less keeled scales on the first dorsal scale rows, always keeled on the posterior part of the body, (2) supralabials

with large, distinct (not contiguous) white (or nearly so) blotches, and (3) a series of dorsal spots on a faint

dorsolateral stripe or at its level if the stripe is not visible, but A. khasiense has only more or less connected

blotches on the neck, which usually do not reach the level of the dorsolateral stripe and small dorsal dots, whereas

A. inas usually shows a distinct, broad, wavy stripe connected to the anterior part of the dorsolateral stripe, and

large dorsolateral spots. In contrast, only A. boulengeri shows a conspicuous dorsolateral stripe on each side and

thin, horizontal pale streaks (rarely isolated blotches, see Fig. 5 bottom) or, more usually, a continuous stripe on

posterior supralabials, which extends as a stripe on the neck. According to these schemes, A. inas is somewhat

intermediate between A. khasiense and A. boulengeri, in sharing the pattern of supralabials of the former species,

the pattern of the neck of the latter one, and enlarged dorsolateral spots, at least on the anterior part of the body.

The most important diagnostic, salient characters separating A. khasiense, A. boulengeri and A. inas are

summarized in Table 5.

TABLE 5. Summary of main, most common characters separating A. khasiense, A. inas and A. boulengeri. See Table 4

for detailed variation.

This great similarity explains the numerous confusions in the literature between these three species. A.

boulengeri is usually easier to recognize with its peculiar patterns on posterior supralabials and its conspicuous

dorsal stripes. However, we have examined some specimens from Vietnam in which the streaks on posterior

supralabials were nearly separated each from the other and thicker than usual. These specimens may easily be

misidentified as Amphiesma khasiense if other morphological characters are neglected. In this case, the keeling of

the 1st dorsal scale row may confirm the identification.

These species and other species of the Amphiesma khasiense group can be identified with the following key. It

was constructed from Smith (1943), Taylor (1965), Ziegler & Le (2006a), David et al. (2007), and from our data.

1 17 (or 15) scale rows at midbody . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

- 19 scale rows at midbody . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2 No temporal; 5–6 SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma atemporale

- At least 1 anterior temporal; 7–8 (rarely 6) SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma sauteri

3 Dorsal pattern with vertical crossbars or incomplete rings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

- Dorsum not banded or ringed, but ornate with dorsolateral dots or stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

4 Narrow pale crossbars on a deep bluish-black background color; no pale spots on head; 152-169 VEN; 7–8 SL . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma optatum

- Wide pale crossbands anteriorly, body not bluish-black; numerous round spots on head; 179 VEN in the sole known male; 9 SL

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Amphiesma andreae

5 Subcaudals single . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma xenura

- Subcaudals paired . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6 A broad, white stripe from the tip of the snout to the corner of the mouth, extending on the nape and forming a conspicuous

chevron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma leucomystax

- Head pattern not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

7 One white vertical streak before and another vertical white stripe just behind the eye; usually 4 PosOc; more than 115 subcau-

dals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesmoides ornaticeps

- Head pattern not as above; 2 or 3 PosOc; less than 115 subcaudals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8 Each posterior supralabial with a distinct, white or cream blotch or streak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Posterior supralabials not as such, i.e. entirely white or cream, diffusely edged with black, or brown with paler areas . . . . . 11

9 Each posterior supralabial with a round or oval, central cream blotch, not forming a stripe; 1st DSR keeled, throughout or at

least on the posterior part of the body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

Posterior supralabials dark brown or black, each with a median, horizontal and longate, thin cream blotch or streak, usually

united as a narrow stripe, extending on the neck as a continuous stripe; scales of 1st DSR entirely smooth . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma boulengeri

Species Supralabials Neck Body 1st DSR

A. khasiense Isolate blotches Spots or streaks Small dots Keeled (in part)

A inas Isolate blotches Stripe Large dots Smooth throughout

A. boulengeri Streaks or stripe Stripe Broad stripe Keeled throughout



10 Dorsal pattern made of a series of reddish-brown or rusty (cream in preservative) dots or small blotches usually aligned on a

faint dorsolateral stripe; usually isolated pale blotches on the side of the neck, sometimes partly connected by a narrow pale

line, not reaching the dorsolateral stripe (if present) or dorsolateral spots; 21–26 maxillary teeth; frontal 2.0–2.5 times longer

than prefrontals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma khasiense

Dorsal pattern made of a series of relatively large yellow or ochre (cream or yellow in preservative) dorsolateral spots on a

more or less visible dorsolateral stripe, the first two to four of them enlarged and connected by a yellow or grey stripe; usually

on the side of the neck a conspicuous, pale stripe or series of large pale, connected spots, reaching the dorsolateral stripe; 19–

21 maxillary teeth; frontal 1.7–1.9 times longer than prefrontals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Amphiesma inas

11 Supralabials entirely white or cream, diffusely edged with black; present in China and Vietnam . . . . . . . . . . . . . . . . . . . . . . 12

- Supralabials brown, heavily dotted with brown, or brown with irregular paler areas; endemic to Sumatra . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma kerinciense

12 Dorsal pattern variegated or reticulated with darker and lighter diffuse blotches; dorsal scales smooth or weakly keeled. . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma johannis

- Dorsal pattern not as above, with dorsolateral stripes; dorsal scales distinctly keeled. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

13 130–142 VEN; 1st DSR smooth; a broad pale blotch on the nape, not touching the posteriormost supralabial . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Amphiesma popei

- 145–172 VEN; 1st DSR keeled; a yellow streak on the nape starting from the posteriormost SL; dorsal scales distinctly keeled

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiesma craspedogaster

Conclusion

If we consider the three species present on the mainland, the morphological characters suggest that A. khasiense

and A. inas are more closely related each with the other than with A. boulengeri. We may consider that we are in

presence of a western complex, including A. khasiense + A. inas, and one eastern, Indo-Chinese species or complex

with A. boulengeri. A. khasiense is mostly present in the eastern and southern parts of the Indo-Himalayan Region.

This species enters northern Thailand and Laos through mountain ranges at the southern fringe of the Indo-

Himalayan system. Quite interestingly, Pauwels et al. (2009) noted that Amphiesma khasiense occurs in Thailand

in three parallel mountain ranges, i.e. from West to East: Thanon Thongchai Range (Doi Inthanon), Khun Tan

Range (Ban Pa Miang Mae Hang) and Phetchabun Range (Phu Luang). This distribution seems to indicate a

penetration from north to south through mountain ranges as in Thailand, A. khasiense has been recorded only high

elevation stations. According to this zoogeographical pattern, and according to the occurrence of A. khasiense in

northern Laos, the occurrence of this species can be expected with little doubt in Phi Phan Nam Range and Louang

Prabang Range, parallel to and between Thanon Thongchai Range and Phetchabun Range. Pauwels et al. (2009)

suggested the occurrence of A. khasiense southwards up to Kanchanaburi Province. Mountains of northern and

western Thailand may act as a corridor allowing the penetration southwards of Indo-Himalayan species. A similar

case is found in, for example, Xenochrophis piscator (Schneider, 1799), which has a similar distribution pattern in

Thailand and northern Laos (Vogel & David, 2012). This scheme may suggest that A. inas, now present to the

mountain ranges of southern Thailand and West Malaysia, evolved from specimens of A. khasiense, or an ancestor

once continuously widespread in the long mountain range edging the Isthmus of Kra. The zoogeography along the

Isthums of Kra has been extensively discussed by Pauwels et al. (2003).

In contrast A. boulengeri is typically an Indochinese species. It is widespread in all mountain ranges of the

eastern and central parts of the Indochinese Peninsula. A. boulengeri may be regarded as a widespread Chinese

species which reached southern parts of the Indochinese Peninsula when the climate of this part of Asia was more

propice. Palaeobiogeography of the region, although imperfectly known, may provide some cues. Heaney (1991)

presented an hypothetical analysis on the evolution of the climate in south-east Asia. During the Pleistocene period

of the Quaternary, south-east Asia was repeatedly affected by a global lowering of sea level, cooler temperatures

and both a higher seasonality and decrease in the volume of rainfalls (see Pauwels et al. [2003] for a more complete

discussion and additional references). It may be suggested that Amphiesma boulengeri was once widespread at all

elevations when the climate of the Indochinese Peninsula was wet and favourable then retreated in elevated places,

wetter than the adjacent lowlands. A similar example may be found in Trimeresurus vogeli David, Vidal &

Pauwels, 2001, closely related to the Chinese species Trimeresurus stejnegeri Schmidt 1925. T. vogeli inhabits only

isolated wet hills or mountain ranges located along the eastern and southern fringes of the extensive lowlands

which ranges covering eastern Thailand, Cambodia, southern Laos and Vietnam.

This paper is based only on morphological characters. Although these widely confused species are

homogeneous in scalation and pattern, the status of the populations here referred to A. boulengeri of the Cardamom



Mountains is not fully solved. A molecular phylogeny based on populations covering the whole range of A.

boulengeri is necessary for a better understanding of relations within this species. In the same way, molecular

analyses would provide cues of the relationships between A. khasiense and A. inas. Lastly, a revision of the genus

Amphiesma, as currently conceived, is badly necessary.

Acknowledgements

We are grateful to Ke Jiang (Kunming Institute of Zoology, Kunming, People’s Republic of China), Mian Hou

(Sichuan Normal University, People’s Republic of China) and Olivier S. G. Pauwels (Institut Royal des Sciences

Naturelles de Delgique, Brussels, Belgium) for their support, comments or comparative pictures. We also want to

thank the following photographers for their help in providing us with their rare pictures: Michael Cota (Natural

History Museum, National Science Museum, Thailand, Pathum Thani, Thailand), Lee Grismer (La Sierra

University, Riverside, USA), Sjon Hauser (Chiang Mai, Thailand), Ashok Kumar Malik (Bangalore, India), Kevin

Messenger (Huntsville, USA), and Truong Quang Nguyen (IEBR, Hanoi Vietnam).

It is also a pleasure to thank the following persons who made this review possible in sending us, 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, New York, USA)Colin J. McCarthy and

Patrick Campbell (BMNH, London, UK), Jens V. Vindum and Alan E. Leviton (CAS, San Francisco, USA),

Yuezhao Wang, Xiaomao Zeng, Jiatang Li and Ermi Zhao (CIB, Chengdu, China), the late Jarujin Nabhitabhata

(CTNRC, Thailand National History Museum, Pathum Thani, Thailand), Harold Voris and Alan Resetar, (FMNH,

Chicago, USA), Truong Quang Nguyen, Canh Xuan Le and Dang Xuan Nguyen (IEBR, Hanoi Vietnam), Michael

W. Lau (KFBG, Hong Kong SAR, People’s Republic of China), Kirati Kunya (KZM, Korat Zoo, Muang District,

Nakhon Ratchasima, Thailand), Lee L. Grismer (LSUHC, Riverside, USA), Alain Dubois, Ivan Ineich et

Annemarie Ohler (MNHN, Paris, France), Meredith J. Mahoney and David Wake (MVZ, Berkeley, USA)Irvan

Sidik (MZB, Bogor, Indonesia), Bryan L. Stuart (NCSM, Raleigh, USA), Lawan Chanhome (QSMI, Bangkok,

Thailand), Jianqiang Cen (SMNH, Shanghai, People’s Republic of China), Nikolai L. Orlov (ZISP, St. Petersburg,

Russia), Mark-Oliver Roedel and Frank Tillack (ZMB, Berlin, Germany), Kelvin P. P. Lim (ZRC, Singapore), and

Channakesava Murphy (ZSI, Kolkata, India).

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APPENDIX. Additional examined specimens.

Amphiesma atemporale (n = 7). Vietnam. Vinh Phu Province. MNHN 1935.0077–78, MNHN 1935.0450, MNHN 1938.0120, MNHN 1958.0461–62, “Tam Dao”, now Tam Dao Hill Station. – No locality. MNHN 1938.0119, “Indochine”, no precise locality.

Amphiesma craspedogaster (n = 11). People’s Republic of China. Fujian Province. BMNH 1910.9.6.2, “South Fokien”; BMNH 1946.1.12.63–65, “Kuatun, N.W. Fokien”, now Guadun, Wuyi Shan, Chongan County; SMNH 2757, SMNH 3592–3594, Congan Shangang. – Guizhou Province. MNHN 1912.0324-26, no locality.

Amphiesma johannis (n = 2). People’s Republic of China. MNHN 1912.0272–73, “Chine occidentale et Marches thibétaines”.

Amphiesma modestum (n = 9). India. Meghalaya. BMNH 76.2.16.1-2, “Cherra Punji, Khasi Hills”, now Cherrapunji; BMNH 1946.1.13.40-41, Khasi Hills. – Myanmar. Kachin State. BMNH 1925.4.2.16, “Hutong, Bhamo District”; BMNH 1925.9.17.2, BMNH 1925.12.22.22-23, “Huton, Kachin Hills”. Kayah State. MNHN 1893.0400, “Mts Carin, 1200-1300 m”, now Mts. Karen.

Amphiesma popei (n = 2). – People’s Republic of China. Guangxi Zhuang Autonomous Region. KFBG 4.VIII.1999, Dawangling. - Vietnam. Vinh Phuc Province. ZISP N-155, Tam Dao.

Amphiesma sauteri (n = 8). – People’s Republic of China. Guizhou Province. MNHN 1912.0327–0328, “Kouy-Tchéou”, no locality. - Hainan Island. KFBG 27.V.1999, Dialuo Shan. – Vietnam. Vinh Phu Province. MNHN 1908.0011, MNHN 1935.0066–0067, MNHN 1935.0454–0455, “Tam Dao, Tonkin”, now Tam Dao Hill Station.

Amphiesma sanguineum (n = 7). Federation of Malaysia. West Malaysia. State of Pahang. ZRC 2.4034, Cameron Highlands, 5500 ft.; ZRC 2.4035, Renglet, Cameron Highlands, 3000 ft.; ZRC 2.4036, ZRC 2.4039, Cameron Highlands; ZRC 2.4038, Telon Valley, Cameron Highlands. State of Selangor. ZRC 2.4037, Gombak Valley; ZRC 2.4040, Dusun Wan, 2000 ft.

Amphiesma xenura (n = 1). Myanmar. Rakhine State. CAS 220550, Daung stream, Gwa Township, 17°35.051N-94°40.689E.


Documents

Genus Amphiesma Dumeril, Bibron & Dumeril, 1854