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REDISCOVERY AND REDESCRIPTION OF TWO RARE SNAKE SPECIES:
Oligodon lacroixi ANGEL ET BOURRET, 1933 AND Maculophis bellus chapaensis
(BOURRET, 1934) [SQUAMATA: OPHIDIA: COLUBRIDAE]
FROM FANSIPAN MOUNTAINS, NORTHERN VIETNAM
Nikolai L. Orlov,1Sergei A. Ryabov,2Tao Thien Nguyen,3and Truong Quang Nguyen4
Submitted May 15, 2010.
We redescribe Oligodon lacroixi and Maculophis bellus chapaensis, two rare snakes which have not been re-
ported from Vietnam since the original descriptions in 1930s, based on three voucher specimens recently col-
lected from Fansipan Mountains, northern Vietnam. The list of colubrid snakes from Fansipan Mountains is also
provided.
Keywords: Oligodon lacroixi;Maculophis bellus chapaensis; colubrids; rare snakes; Fansipan; Vietnam.
INTRODUCTION
The snake fauna of the tropical rain forest is charac-
terized not only by the species diversity but also by the
extremely rare repetition of the records in a number of
species. For example, in the rain- and monsoon forests
of Vietnam many taxa of snakes are known only from
the type series or a single specimen (Orlov et al., 2003).
Both Oligodon lacroixi and Maculophis bellus chapaen-
sis were originally described based on the type speci-
mens from Vietnam. However, since the descriptions by
Angel and Bourret (1933) and by Bourret (1934), no
additional voucher specimens of above taxa had been
reported from this country. Based on the recent collec-
tion of barely known snakes from Fansipan Mountains,
northern Vietnam, we herein expand the species de-
scription.
Earlier they were not found despite of intensive zoo-
logical research. Probably it could be explained that it is
a phenomenon of cyclic changes of population density
with unknown duration of such a cycle. That is possible
that increasing intensity of field study in this region
leads to positive results and allow repeating records of
rare snakes.
MATERIAL AND METHODS
Field work on the herpetofauna was conducted in Sa
Pa, Lao Cai Province in 2009 and 2010. Specimens re-
ferred to this paper are deposited in the collections of the
Institute of Ecology and Biological Resources (IEBR),
Vietnam; Kunming Institute of Zoology (KIZ), China;
Museum National d’Histoire Naturelle (MNHN),
France, and Zoological Institute, St. Petersburg (ZISP),
Russia.
All measurements were taken with a caliper to the
nearest 0.1 mm. Morphological characters were exam-
ined with a binocular microscope Leica MZ 8 and pho-
tographs were taken with a camera Nikon D700 with
lens Nikkor-micro 105 mm.
RESULTS
Oligodon lacroixi Angel et Bourret, 1933 (Fig. 1)
Oligodon lacroixi F. Angel et R. Bourret, 1933, Bull.
Soc. Zool. France, 58:138.
Type locality. Chapa (= Sa Pa, Lao Cai Prov.),
Tong-king (= northern Vietnam); 1500 m.
Types. Lectotype MNHN 1772-2 (1933-1) (Fig. 2);
Paralectotype MNHN 1772-3 (1933-1a).
Distribution. Beside the type series as mentioned
above, few voucher specimens of this species had been
reported: five specimens from the type locality in Viet-
1026-2296/2010/1704-0310 © 2010 Folium Publishing Company
Russian Journal of Herpetology Vol. 17, No. 4, 2010, pp. 310 – 322
1Zoological Institute, Russian Academy of Sciences, Universitet-
skaya nab., 1, St. Petersburg 199034, Russia;
e-mail: azemiops@zin.ru
2Tula Exotarium, Oktyabr’skaya ul., 26, Tula 300002, Russia.
3Vietnam National Museum of Nature, 18 Hoang Quoc Viet Road,
Cau Giay, Hanoi, Vietnam; e-mail: nguyenthientao@gmail.com
4Institute of Ecology and Biological Resources, 18 Hoang Quoc Viet
Road, Hanoi, Vietnam; e-mail: E-mail: nqt2@yahoo.com;
Zoologisches Forschungsmuseum Alexander Koenig, Adenauer-
allee 160, D-53113 Bonn, Germany.
nam, deposited in the collection of the Museum National
d’Histoire Naturelle MNHN 1935-4, 1938-137, 1938-
138, 1958-464, 1958-464a) (Bourret, 1934, 1935, 1936,
1937, 1939); a female specimen collected in June 1977
in vicinity of Mengzi, Honghe prefecture, Yunnan Prov-
ince, China, deposited in the collection of the Kunming
Institute of Zoology (KIZ 0331) (He and Yang, 1979;
Zhao and Adler, 1993; Zhao et al., 1998; Zhao, 2006)
(Fig. 3); and another female from vicinity of Panzhihua
in the south of Sichuan Province (Zhao, 2006). Thus re-
liable records are known from border regions between
Vietnam and China (within north of Lao Cai province
and south-east Yunnan province) and southernmost re-
gion of Sichuan Province (Fig. 4).
Specimens examined. ZISP 25956 (adult female),
collected by Sergei Ryabov and Nguyen Thien Tao
in June 2010, from Sa Pa, Lao Cai Province, Vietnam,
altitude 1500 m a.s.l. (Fig. 1); Bourret’s collection at
MNHN: 1772-2 (1933-1); 1772-3 (1933-1a); KIZ 0331
(Figs. 2 and 3).
Description of adult female (ZISP 25956). Total
length 805 mm; snout-vent length 705 mm; tail length
100 mm; ventrals 163; subcaudals 34 pairs; anal plate
divided; dorsal scale rows at body [1, at the level of the
15th ventral shield from the head; 2, at midbody; 3, at the
level of the 15th ventral shield from the anal plate]: 1 —
15, 2 — 15, 3 — 15; supralabials 5 (second and third in
contact with eye); infralabials 6; rostral distinctly visible
from above, turned far back; loreal absent; supraocular
single; preocular single; postoculars 2; nasal single,
large, and in contact with rostral, first supralabial and
prefrontal; nostril split posterior margin of nasal; pre-
frontals large, wider than long; supraocular single, very
large, a bit shorter than length of frontal; frontal large,
hexagonal, longer than wide; parietals large, as wide as
frontal; temporals 3, 1 in anterior and 2 in posterior;
internasal absent; mental small, in shape of isosceles tri-
angle; chin shields in 2 pairs, large, two anterior pair
large, in contact with first to fourth infralabials, poste-
rior pair about a half the anterior, in contact with fourth
infralabial.
Coloration in life. Body dark brown above, irides-
cent, with four indistinct black longitudinal stripes,
median pair bordering vertebral series of scales; a verte-
bral series of small orange, rounded, black-edged spots.
Belly orange with alternate transverse black bars, in
Oligodon lacroixi and Maculophis bellus chapaensis from Northern Vietnam 311
Fig. 1. Oligodon lacroixi ZISP 25956, general view.
width of 1 (rarely 2) ventral scales, and its length about
one half of ventral scale. Head dark brown above with 3
light orange, oblique, in Ë-shape markings: one behind
rostral, one behind eyes, and another one in posterior.
Taxonomic comments. Pope (1935) had some
doubts that O. lacroixi is a distinct species and supposed
that it is conspecific with burmanese species O. hampto-
ni Boulenger, 1918. Smith (1943) also noted that it is
312 Nikolai L. Orlov et al.
Fig. 2. Lectotype of Oligodon lacroixi [MNHN 1772-2 (1933-1)].
Fig. 3. Oligodon lacroixi (KIZ 0331) Mengzi, Honghe prefecture, Yunnan province, China.
“like hamptoni in general scalation,” but considered it as
distinct species on the base of original description. How-
ever, O. hamptoni can be distinguished from O. lacroixi
by having a small loreal, temporals 1 + 1, postocular 1,
TL 590 mm, back with a broad yellow vertebral stripe
(vs. loreal absent; temporals 1 + 2, postoculars 2; TL
800 mm; back with a series of orange vertebral spots)
(Bourret, 1936; Smith, 1943; Zhao, 2006; personal ob-
servation). Our specimen has no differences from origi-
nal description.
Maculophis bellus (Stanley, 1917)
Coronella bella A. Stanley, 1917, Jour. North-China
Br. Roy. Asiatic Soc., Shanghai, 47:84.
Maculophis bellus — F. T. Burbrink and R. Lawson,
2007, Mol. Phylogen. Evol., 43(1): 173 – 189.
Distribution. Distribution range is disjunctive: min-
imum six isolated parts are known (1, Yunnan –
Myanmar; 2, Fujian; 3, Guangxi; 4, Hunan; 5, Ha Tinh;
6, north and northwest Vietnam). This species has been
reported from southern mountain regions of China (Yun-
nan, Guangxi, Hunan, and Fujian provinces), northern
and northwest Vietnam, northern Myanmar and proba-
bly northern Laos at elevation between 1000 – 3000 m
a.s.l. (Pope, 1935 as Coronella bella; Bourret, 1934,
1936 as Elaphe leonardi leonardi,E. l. chapaensis;
Smith, 1943 as Elaphe leonardi leonardi,E. l. chapaen-
sis; Deuve, 1970 as Elaphe leonardi; Ding and Zheng,
1974 as Oligodon bella; Welch, 1988 as Elaphe leonardi
leonardi,E. l. chapaensis; Zhao and Adler, 1993 as Oli-
godon bellus; Schulz, 1996 as Elaphe leonardi leonardi,
E. l. chapaensis; Zhao et al., 1998 as Oligodon bellus;
Schulz et al., 2000 as Elaphe bella; Helfenberger, 2001
as Elaphe bella; Jiang et al., 2006 as Oligodon bellus;
Zhao, 2006 as Oligodon bellus; Burbrink and Lowson,
2007 as Maculophis bellus; Nguyen, 2007 as Elaphe
leonardi; Orlov and Ryabov, 2009 (cover) as Macul-
ophis bellus chapaensis; Nguyen et al., 2009 as Elaphe
bella leonardi,E. b. chapaensis) (Fig. 4). Complicated
mountain relief on the border of several East Asian
countries does not allow having exact ideas about distri-
bution range of this species. Rare records in the territory
from Fujian province in the east to northern Myanmar in
the west and from north-west of Yunnan province to Lao
Cai province at the altitude about 1000 – 3000 m a.s.l.,
are very sporadically distributed on the map of mountain
East Aia. It is explained mainly by very scarce survey of
this vast region.
This species includes two subspecies:
Maculophis bellus bellus (Stanley, 1917)
Coronella bella A. Stanley, 1917, J. N.-China Branch
Roy. Asiatic Soc., Shanghai, 47:84.
Type locality. Kuatun (= Guadun) Hills, northwest
Fokien (= Fujian Prov.), China.
Types. Fujian University, Department of Biology,
Shanghai, Guangdong Province, China.
Distribution. This subspecies is distributed in
southern mountain regions of China (Yunnan, Guangxi
Zhuang Autonomous Region, Hunan and Fujian prov-
inces, Guangxi Zhuang Autonomous Region) and north-
ern Myanmar (Sinlum Kaba, Kambaiti and Patsarlam-
dam, Kachin State of Myanmar) (Pope, 1935; Bourret,
1934, 1936; Smith, 1943; Deuve, 1970; Ding and
Zheng, 1974; Zhao and Adler, 1993; Schulz, 1996; Zhao
et al., 1998; Jiang et al., 2006; Zhao, 2006; Nguyen,
2007; Nguyen et al., 2009). In 2010 we got additional in-
formation about new records from Guangxi Zhuang Au-
tonomous Region and West Mangshan, Hunan Province
and photographs of M. b. bellus from Chinese and Dutch
colleagues (Wang Zhimin, Hang Yang and Gidi van de
Belt).
Oligodon lacroixi and Maculophis bellus chapaensis from Northern Vietnam 313
Fig. 4. Distribution and type locality of Oligodon lacroixi,Macul-
ophis bellus bellus, and Maculophis bellus chapaensis:, distribu-
tion of Oligodon lacroixi (Sa Pa, Lao Cai Province of Vietnam; Meng-
zi, Yunnan Province and Panzhihua, Sichuan Province of China);
, distribution of Maculophis bellus bellus (Kambaiti, Patsarlamdam
and Sinlum Kaba, Kachin State of Myanmar; Yunnan, Guangxi
Zhuang Autonomous Region, Hunan and Fujian Provinces of China);
, distribution of Maculophis bellus chapaensis (Lao Cai, Lang Son,
Bac Thai, Vinh Phuc, and north-west of Ha Tinh Provinces of Viet-
nam); T1, type locality of Maculophis bellus chapaensis; T2, type lo-
cality of Oligodon lacroixi; T3, type locality of Maculophis bellus
bellus;, North Laos (locality Maculophis bellus chapaensis sug-
gested by Deuve, 1961).
314 Nikolai L. Orlov et al.
Fig. 5. Maculophis bellus bellus (KIZ 75 II 0180), dorsal view.
Fig. 7. Maculophis bellus bellus (KIZ 78 I 001), dorsal view. Fig. 8. Maculophis bellus bellus (KIZ 78 I 001), ventral view.
Fig. 6. Maculophis bellus bellus (KIZ 75 II 0180), ventral view.
Specimens examined. Elaphe leonardi (= Macul-
ophis bellus –“Maculophis bellus bellus”): KIZ 75 II
180, KIZ 78 II 002; 78 II 001; 75 II 200; 75 II 0014;
75 II 0136; 75 II O147 — Muodaohe and Bijiang (= Zhi-
zi Luo), Jingdong (Wuliang Mountain), Yunnan, China;
1900 – 2400 m (Figs.5–9).
Taxonomic comments. Some differences in color
patterns are recorded between individuals of “East
China” populations (Hunan, Guangxi and Fujian prov-
inces) and “western” populations from Myanmar and
Yunnan province, China (Figs.5–11). Second subspe-
cies, Maculophis bellus chapaensis (Bourret, 1934) is
known from northern Vietnam and probably from north-
ern Laos.
Maculophis bellus chapaensis (Bourret, 1934)
Elaphe leonardi chapaensis R. Bourret, 1934, Bull.
Gén. Instr. Publique, Hanoi, 1934(7): 7.
Type locality. Chapa, Tongking (now Sa Pa, Lao
Cai Province, northern Vietnam).
Types. Lectotype MNHN 1938-01322 (1164-1)
(Fig. 12); paralectotypes: MNHN 1958-0467, 1958-
0468.
Distribution. This subspecies previously known
only from Vietnam: Sa Pa (Fan Si Pan Mountain), Lao
Cai Province (e.g., Bourret, 1934, 1935, 1936, 1937;
Schulz, 1996; Nguyen and Ho, 1996; Nguyen et al.,
2009; Orlov and Ryabov, 2009 (cover RJH). Additional
record from Lang Son, Bac Thai, Vinh Phuc, and
north-west of Ha Tinh provinces were reported by Orlov
et al. (2000), and followed by Nguyen et al. (2009) with-
out voucher museum specimens and based only on ob-
servations, literature and hunter communications
(Nguyen and Ho, 1996). Seven specimens were regis-
tered in 2008 – 2010 in the nature; 2 specimens (adult
females) were collected in May 2009 and June 2010 in
vicinity of Sa Pa, altitude about 1900 m a.s.l.: ZISP
25954 and ZISP 25955 (Fig. 5). Deuve (1970) suggests
the presence of this species in northern Laos.
Specimens examined. Elaphe leonardi chapaensis
(= Maculophis bellus chapaensis): Bourret’s collection
at lectotype MNHN 1164-1 (1938-01322) and paralecto-
types MNHN 1958-0467, 1958-0468; Maculophis bel-
lus chapaensis: ZISP 25954 and ZISP 25955 (two adult
females) — Sa Pa (Fan Si Pan Mountain), Lao Cai prov-
ince (Fig. 4).
Oligodon lacroixi and Maculophis bellus chapaensis from Northern Vietnam 315
Fig. 9. Maculophis bellus bellus (KIZ 75 II 0014).
Description of two adult females (ZISP 25954 and
ZISP 25955). (Figs. 13 – 15). Total length — 931 and
836 mm; snout-vent length — 796 and 713 mm; tail
length from vent to tip — 135 and 123 mm; ventrals —
211 and 209; subcaudals — 54 and 53 pairs; anal plate
1 – 1 and 1 – 1 (divided); dorsal scale rows at body [1, at
the level of the 15th ventral shield from the head; 2, at
midbody; 3, at the level of the 15th ventral shield from
the anal plate]: 19 – 19 – 17 and 19 – 19 – 17; supralabi-
als 6 – 6 and 7 – 7 (third and fourth in contact with eye);
infralabials 9 – 9 and 9 – 9; rostral very large, wide and
high, clearly visible from above; no loreal; supraocu-
lar — 1 and 1; preocular — 1 and 1; postoculars — 2 and
2; nasal paired, large nostril cut between anterior and
posterior nasal, anterior nasal in contact with rostral,
internasal and first supralabial, posterior nasal in contact
with prefrontal, internasal, preocular, first and second
supralabials; paired internasals considerably wide than
longer; paired prefrontals in their shape and size almost
similar with internasals, their width greater that length;
mental very large, its width slightly greater than its
length; single supraoculars, very large, their length
slightly less that length of frontal; frontal large, hexago-
nal, its length greater than width; large, wide parietals in
their length greater than frontal anterior temporal —
2–3and2–2,posterior temporal—2–2and2–2.
316 Nikolai L. Orlov et al.
Fig. 10. Maculophis bellus bellus from Western Mangshan, Hunan prov., China. Photo by Gidi van de Belt.
Fig. 11. Maculophis bellus bellus from Guangxi Zhuang Autonomus
Region. Photo by Wang Zhimin and Hang Yang.
Oligodon lacroixi and Maculophis bellus chapaensis from Northern Vietnam 317
Fig. 13. Maculophis bellus chapaensis (ZISP 25954) from Sa Pa (Fan Si Pan Mountain), Lao Cai province, Vietnam, dorsal view.
Fig. 12. Lectotype MNHN 1164-1 (1938-01322) of Maculophis bellus chapaensis.
318 Nikolai L. Orlov et al.
Fig. 14. Maculophis bellus chapaensis (ZISP 25954) from Sa Pa (Fan Si Pan Mountain), Lao Cai province, Vietnam, ventral view.
Fig. 15. Maculophis bellus chapaensis ZISP 25955 from Sa Pa (Fan Si Pan Mountain), Lao Cai province, Vietnam, general view and head.
Coloration in life. Body of adults reddish brown
above (bright red in juveniles) with 27–29 transverse
olive, black-edged bands (brightly yellow bands in
black edging in juvenile snakes), some bands in Ë-shape
on lower lateral zone and black edge contiguous to
ventral scales usually broken into black spots; rostral
and loreal areas yellow; dorsal head with an yellow
marking, black-edged, in V-shape, from above eye to
neck, frontal usually with yellow streak medially
(Figs. 12, 15, and 16).
Taxonomic comments. Until now taxonomic status
of forms “bellus” and “chapaensis” is still questionable.
Burbrink and Lawson (2007) in their study of phylogeny
of colubrids validate the taxon Maculophis. They used
in molecular analysis only tissues of specimens from un-
identified province of China; there were no DNA materi-
als of “chapaensis” from Vietnam up to 2007. They con-
clude that the oldest available genus that should be ap-
plied to the monophyletic group of E. frenata and
E. prasina is Rhadinophis (Williams et Wallach, 1989).
According to Burbrink and Lawson: «The other taxon,
E. bella, has no available genus and retaining Elaphe for
that taxon makes the genus paraphyletic. Given that
E. bella does not have priority for the genus Elaphe,we
suggest the name Maculophis, which, when translated,
means spotted (maculos) snake (ophis) and is in refer-
ence to the blotched dorsum. Thus, this species would
now be Maculophis bellus. We are aware, however, that
this does create the unfortunate situation of proposing a
monotypic genus». There are slight differences between
Oligodon lacroixi and Maculophis bellus chapaensis from Northern Vietnam 319
Fig. 16. Newborn of Maculophis bellus chapaensis.
Fig. 17. Habitat of Maculophis bellus chapaensis in the northeastern slope of Fan Si Pan Mountain (Sa Pa, Lao Cai province, Vietnam).
“bella” and “chapaensis” in pholidosis and coloration:
anterior temporals in M. b. bellus —1–1,inM. b. cha-
paensis — 2 – 2, 2 – 3; across the body (without tail) in
M. b. bellus — 39 – 46 bands, in M. b. chapaensis —
27 – 29 bands. Color pattern differences are clearly
shown by Schulz (1996).
Notes. Oligodon lacroixi and Maculophis bellus
chapaensis are found at the altitude 1500 – 1900 m a.s.l.
on the slope of mount Fan Si Pan (Fig. 17) located in the
Hoang Lien mountains, which are an extension of the
Ailao Shan mountains in China, the south-eastern extent
of the Himalayan chain (Tordoff et al. 1999). Fan Si Pan
mount is inhabited by unique and diverse herpetofauna
including snakes of family Colubridae sensu lato.Were
-
corded 65 species of 32 genera from 1987 at the altitude
from 400 to 3000 m a.s.l.
List of ophidiofauna of Fan Si Pan Mountain in-
cludes Eastern Himalayan-Tonkin and South-Indochina
elements in their composition.
LIST OF COLUBRIDAE OF FAN SI PAN
MOUNTAIN:
Colubrinae Cope, 1898
Ahaetulla Link, 1807
Ahaetulla prasina (Reinhardt, 1827)
Boiga Fitzinger, 1826
Boiga cyanea (Duméril, Bibron et Duméril, 1854)
Boiga guangxiensis Wien, 1998
Boiga multomaculata (Boie, 1827)
Cyclophiops Boulenger, 1888
Cyclophiops major (Günther, 1858)
Cyclophiops multicinctus (Roux, 1907)
Coelognathus Fitzinger, 1843
Coelognathus radiatus (Schlegel, 1837)
Dendrelaphis Boulenger, 1890
Dendrelaphis ngansonensis (Bourret, 1935)
Dendrelaphis pictus (Gmelin, 1789)
Dinodon Duméril et Bibron, 1853
Dinodon meridionale Bourret, 1935
Dinodon rufozonatum (Cantor, 1842)
Dinodon septentrionalis (Günther, 1875)
Elaphe Fitzinger, 1833
Elaphe carinata ornitophaga Bourret, 1936
Euprepiophis Fitzinger, 1843
Euprepiophis mandarinus (Cantor, 1842)
Liopeltis Fitzinger, 1843
Liopeltis frenatus (Günther, 1858)
Lycodon Boie, 1826
Lycodon fasciatus (Anderson, 1879)
Lycodon ruhstrati (Fischer, 1886)
Lycodon subcinctus Boie, 1827
Maculophis Burbrink et Lowson, 2007
Maculophis bellus chapaensis (Bourret, 1934)
Oligodon Boie, 1827
Oligodon albocinctus (Cantor, 1839)
Oligodon chinensis (Günther, 1888)
Oligodon cinereus (Günther, 1864)
Oligodon cyclurus (Cantor, 1839)
Oligodon eberhardti Pellegrin, 1910
Oligodon lacroixi Angel et Bourret, 1933
Oligodon taeniatus (Günther, 1861)
Oreocryptophis Uttiger, Schätti et Helfenberger, 2005
Oreocryptophis porphyraceus vaillanti (Sauvagi, 1876)
Orthriophis Uttiger, Schätti Schmidt,
Ruf et Ziswiler, 2002
Orthriophis taeniurus yunnanensis (Anderson, 1879)
Ptyas Fitzinger, 1843
Ptyas korros (Schlegel, 1837)
Ptyas mucosa (Linnaeus, 1758)
Rhadinophis Williams et Wallach, 1989
Rhadinophis frenatus (Gray, 1853)
Rhadinophis prasinus (Blyth, 1854)
Sibynophis Fitzinger, 1843
Sibynophis chinensis (Günther, 1889)
Sibynophis collaris (Gray, 1853)
Zaocys Cope, 1861
Zaocys carinatus (Günther, 1858)
Zaocys nigromarginatus (Blyth, 1854)
Subfamily Calamariinae
Calamaria Boie, 1826
Calamaria pavimentata Duméril, Bibron et Duméril,
1854
Calamaria septentrionalis Boulenger, 1890
Natricinae Bonaparte, 1840
Amphiesma Duméril, Bibron and Duméril, 1854
Amphiesma bitaeniata (Wall, 1925)
Amphiesma boulengeri (Gressitt, 1937)
Amphiesma khasiensis (Boulenger, 1890)
Amphiesma modesta (Günther, 1875)
Amphiesma optata (Hu et Zhao, 1966)
Amphiesma parallela (Boulenger, 1890)
Amphiesma stolata (Linnaeus, 1758)
320 Nikolai L. Orlov et al.
Opisthotropis Günther, 1872
Opisthotropis jacobi Angel et Bourret, 1933
Pararhabdophis Bourret, 1934
Pararhabdophis chapaensis Bourret, 1934
Rhabdophis Fitzinger, 1843
Rhabdophis nuchalis (Boulenger, 1891)
Rhabdophis subminiatus (Schlegel, 1837)
Sinonatrix Rossman et Eberle, 1977
Sinonatrix aequifasciata (Barbour, 1908)
Sinonatrix percarinata (Boulenger, 1899)
Psammodynastes Günther, 1858
Psammodynastes pulverulentus (Boie, 1827)
Xenochrophis Günther, 1864
Xenochrophis flavipunctatus (Hallowell, 1860)
Pseudoxenodontinae McDowell, 1987
Plagiopholis Boulenger, 1893
Plagiopholis delacouri Angel 1929
Plagiopholis nuchalis (Boulenger, 1893)
Plagiopholis styani (Boulenger, 1899)
Pseudoxenodon Boulenger, 1890
Pseudoxenodon bambusicola Vogt, 1922
Pseudoxenodon macrops (Blyth, 1854)
Pareatinae Romer, 1956
Pareas Wagler, 1830
Pareas carinatus (Boie, 1828)
Pareas hamptoni (Boulenger, 1905)
Pareas macularius Theobald, 1868
Pareas margaritophorus (Jan, 1866)
Pareas monticola (Cantor, 1839)
Xenodermatinae Cope, 1900
Achalinus Peters, 1869
Achalinus spinalis Peters, 1869
Homalopsinae Bonaparte, 1845
Enhydris Sonnini et Latreille, 1802
Enhydris plumbea (Boie, 1827)
Acknowledgments. We are very grateful to Vietnam’s In-
stitute of Ecology and Biological Resources (IEBR) for the or-
ganization of the fieldwork in Vietnam, Director of IEBR,
Prof. Le Xuan Canh and Director of Dept. of Vertebrate Zool-
ogy, Dr. Nguyen Xuan Dang.
We are indebted to Annemarie Ohler and Patrick David
(MNHN), Ho Thu Cuc (IEBR) and Rao Ding-qi (KIZ) for per-
mitting us to examine snake specimens. Thanks a lot to Wang
Zhimin, Hang Yang (China) and Gidi van de Belt (Holland)
for providing photographs to us. This research was partially
supported by the Project “Fauna of Vietnam,” Grant NSH
1647.2003 Volkswagen and the German Academic Exchange
Service (DAAD), Grant ScSh 4724.2010.4 and RFBR
08-04-00041.
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