Russian Journal of Marine Biology, Vol. 30, No. 6, 2004, pp. 387–394.

Original Russian Text Copyright © 2004 by Biologiya Morya, Kharin.

INVERTEBRATE
ZOOLOGY

Review of Sea Snakes of the Genus Hydrophis

sensu stricto (Serpentes: Hydrophiidae)
V. E. Kharin
Institute of Marine Biology, Far East Division, Russian Academy of Sciences,
Vladivostok, 690041 Russia
e-mail: vkharin@imb.dvo.ru
Received November 14, 2003

Abstract—The results of the genus Hydrophis revision s. str. are presented. The genus is divided into three sub-
genera, including Mediohydrophis subgen. nov. The validity of Hydrophis atriceps and H. (Microcephalophis)
gracilis microcephalus is confirmed. New data on the distribution of H. klossi is given.

Key words: Hydrophis, sea snakes, revision, validity, subspecies, subgenera.

Microcephalophis was first established by Lesson
[23] for the snake Hydrus gracilis Shaw, 1802. There-
after, it was considered in synonymy of the genera Dis-
teira (= Distira) [39, 45] or Hydrophis [2, 3, 11, 15, 44].
Wall [45] resurrected it for two species, H. gracilis and
H. cantoris Gunther, 1864, later Smith [35, 37] sup-
ported him. McDowell [24] again reduced this genus to
synonymy of Hydrophis s. str. on the basis of morpho-
logical and osteological studies; in subsequent works
this fact was either contested [43] or accepted [8, 9, 26].
As a result of the revision of this group [18, 19, 20]
McDowell’s point of view was confirmed, which led to
the division of the genus Hydrophis into five species
groups, which are suggested by us as subgenera
[20, 22]. In the last annotated catalogues of poisonous
snakes [13, 29, 47] the whole polytypic generic group
Hydrophis sensu lato is considered as the monotypic
genus Hydrophis without adequate analysis of the
osteological characteristics. Following McDowell [24]
we shall try to prove the polytypy of this group divided
into five separated genera [19, 22].
MATERIALS AND METHODS
The investigations were based on the collection of
the Zoological Institute of the Russian Academy of Sci-
ences (ZIN) and comprehensive materials of the Pacific
Fisheries Research Center (TINRO-Center). Additional
materials were gained from the British Museum of Nat-
ural History (BMNH), the American Museum of Natu-
ral History (AMNH), and the Berlin Zoological
Museum of Humboldt University. Twelve specimens of
Hydrophis cantoris were studied from the eastern coast
of India (including ZMB 2837, one of the syntypes of
Hydrophis fasciatus); 36 specimens of H. gracilis from
the eastern coast of India, the Gulf of Siam, Indonesia,
Australia, the Gulf of Tonkin, and the southern China
coast; 18 specimens of H. klossi from the Gulf of Siam
and Malaysian coastal waters; 9 specimens of H. mel-
anosoma from Malaysian coast and the north Austra-
lian shelf; 23 specimens of H. obscurus from the east-
ern coast of India and the Andaman Sea; type specimen
of H. vorisi from the southern coast of New Guinea
(AMNH 58869); 16 specimens of H. brookii from the
Gulf of Siam and the coastal waters of Vietnam; 8 spec-
imens of H. macdowelli (including ZIN 19678, 19679,
19680 type series) from the north Australian shelf;
14 specimens of H. atriceps (including ZMB 7436,
paralectotype of Disteira cincinatii) from the Gulf of
Siam, the Philippine coast, Indonesia, and the north
Australian shelf; and 17 specimens of H. fascistus from
the eastern coast of India and the Bay of Bengal. The
data about the geographical distribution was borrowed
from Smith’s [35] and Minton’s [27] works.
In this paper standard measurement methods and the
abbreviations of folidose features were used [17].
SYSTEMATIC PART
Genus Hydrophis Latreille in Sonnini et Latreille,
1802.
Hydrophis Latreille in Sonnini et Latreille, 1802:
193 (type species Hydrophis laticauda Latreille in Son-
nini et Latreille, 1802 = Hydrus fasciatus Schneider,
1799, by subsequent designation—Smith, 1926: 41).
Microcephalophis Lesson in Belanger, 1834: 321
(type species Hydrus gracilis Shaw, 1802, by mono-
typy).
Liopala Gray, 1842: 60 (type species Hydrus graci-
lis Shaw, 1802, by monotypy). Porrecticollis Wall,
1921: 335 (type species Hydrophis obscurus Daudin,
1803, by original designation).

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388 KHARIN

1b, 1c). Generally the palatinum has a triangular prom-

(a) (d) pf inence for the maxillare on the inner side (except
m p
H. macdowelli). The maxillare has corresponding
openings for this prominence. The sphenoideum
widely enters beyond the ventral margin of the cavum
ept epiptericum opening. The quadratum is vertical. The
pt
pr parietale is connected with the praefrontale either
po f totally and separates the frontale from the orbital mar-
gin and from the postorbitale (Figs. 1d, 1e) or incom-
pletely and then the frontale enters partially into the
orbital margin and contacts the postorbitale (Fig. 1f).
The maxillare bears 5–7 maxillary teeth separated by
diastema from poisonous fangs, the palatinum has
6−9 teeth, the pterygoideum has 6–17 teeth, and the
dentale has 11–6 teeth.
(b) (e) The heart is placed slightly forward or behind the
midbody. Hemipenis is covered by strong spines.
The head is small or of moderate size, its scales of a
regular symmetrical shape; eye diameter is equal to the
distance between it and mouth edge and sometimes it is
somewhat less or more than that distance. On infralabi-
als slitlike scales can be present. The first four infrala-
bials contact the mandibular plates. The first pair of
mandibular plates is well developed; they contact each
other. Body scales lay tegularly or almost so, in the pos-
terior part of the body they are hexangular, located side
by side. Body scales, as a rule, are smooth in the ante-
rior part and with the elongate keel or tubercles in the
middle and posterior parts. Ventrals are small, smooth,
(c) (f) and sometimes divided by the elongated groove in the
posterior part.
The genus includes 11 species distributed from the
Persian Gulf and the Arabian Sea to Australia, the Phil-
ippines, and southern China.

KEY TO SUBGENERA AND SPECIES

Fig. 1. Upper jaw apparatus in specimens of the subgenera:
a—Microcephalophis, b—Mediohydrophis, c—Hydrophis
s. str. and the cranium in d—Microcephalophis, e—Medio-
hydrophis, f—Hydrophis s. str. Designations: ept—ectop-
terygoideum, f—frontale, m—maxillare, p—palatinum,
pf—praefrontale, po—postorbitale, pr—parietale, pt—
pterygoideum.
Micromastophis Wall, 1921: errata paste -in the ref-
erence on p. 344 [type species Aturia fasciata [31], by
the original designation].
Diagnosis. The maxillare is not protruded before the
palatinum or is behind it, longer than the ectopterygoi-
deum, its anterior part is embowed so the poisonous
fang lies on a level with the maxillary teeth though it is
significantly longer than any one of them (see Figs. 1a,
OF THE GENUS HYDROPHIS
(1) Ventrals are divided by the elongated groove in
the posterior part of the trunk. The maxillare
protrudes before the palatinum (Fig. 1a). The
parietale is connected with the praefrontale and
separates the frontale from the orbital edge and
from the postorbitale (Fig. 1d). There are no
slitlike scales on the infralabials. (Subgenus
Microcephalophis).........................................2.
Ventrals are smooth in the posterior part of the
trunk. The maxillare does not protrude before
the palatinum (Fig. 1b, 1c). The parietale is con-
nected with the praefrontale either in whole and
separates the frontale from the edge of the orbit
and from the postorbitale or connected incom-
pletely and then the frontale contacts partially
with the orbital margin and with the postorbitale
(Fig. 1e, 1f). Slitlike scales are usually present ....3.
(2) Prefrontal plates contact the third upper labial.
41–48 scales around the midbody, 404–468
ventrals ...........................................H. cantoris.

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 REVIEW OF SEA SNAKES
Prefrontal plates do not contact the third upper
labial. 30–43 scales around the midbody,
220−350 ventrals..............................H. gracilis.
(3) The parietale always connected with the prae-
frontale so that the frontale separated from
orbital edge and the postorbitale (Fig. 1e).
(Subgenera Mediohydrophis nov.)...................4
The parietale is not completely connected with
the praefrontale so that the frontale enters par-
tially into the margin of the orbit and contacts
the postorbitale (Fig. 1f). (Subgenera Hydro-
phis s. str.).......................................................5
(4) Slitlike scales are absent on infralabials. Body
scales smooth or with little keel. 360–430 ven-
trals. The pterygoideum with 7–10 teeth,
12−13 teeth on the dentale...................H. klossi.
Slitlike scales are usually present on infralabials.
Body scales with strong keel. 266–368 ventrals.
The pterygoideum with 9–12 teeth, 13–14 teeth
on the dentale...............................H. melanosoma.
(5) There are 19–23 scales around the neck. The
palatinum always with prominence for the max-
illare.................................................................6
There are more than 23 scales around the neck;
if they number 23, the palatinum is without
prominence for the maxillare...........................8
(6) On infralabials slitlike scales are absent. Body
scales lay almost tegularly in posterior part,
their form is near to hexangular. It is an inhabit-
ant of the South Chinese Sea....... H. parviceps.
Slitlike scales are present. Body scales lay tegu-
larly in the poserior part, the form is not hexan-
gular.................................................................7
(7) 73–74 crossbands around the body and the tail.
It dwells on the southern coast of New
Guinea............................................... H. vorisi.
34–60 crossbands around the body and the tail.
It occurs in the Andaman Sea and on the coast
of East India..................................H. obscurus.
(8) 235–290 ventrals. The palatinum without the prom-
inence for maxillare....................H. macdowelli.
Ventrals always more than 320. The palatinum
has a prominence for the maxillare..................9
(9) The head dark with yellow strip through the
snout and along its sides. 37–45 scales around
the midbody. 328–453 ventrals........ H. brookii.
The head black or dark-olive with yellowish
oval specks on each side. 39–58 scales around
the midbody. 323–514 ventrals......................10.
(10) 27–33 scales around the neck and 49–58 scales
around midbody. 410–514 ventrals (usually
more than 455).............................H. fasciatus.
25–30 scales around the neck and 39–49 scales
around midbody. 323–453 ventrals (usually less
than 366)...........................................H atriceps.
RUSSIAN JOURNAL OF MARINE BIOLOGY Vol. 30
389
Subgenus Microcephalophis Lesson in Belanger,
1834
Type species: Hydrus gracilis Shaw, 1802.
Diagnosis. The maxillare protrudes before the
palatinum (Fig. 1a). The parietale is always connected
with the praefrontale so that the frontale is separated
from the orbital edge and from the postorbitale
(Fig. 1d). In the posterior part of the body the ventrals
are divided in half by an elongated groove.
The subgenus includes two species.
Hydrophis (Microcephalophis) gracilis (Shaw,
1802).
Hydrus gracilis Shaw, 1802: 560 (the type locality
is unknown).
Hydrophis Kaddell-Nagam Boie, 1827: 554 (Bay of
Bengal, the description is based on Russell, 1801:
pl. XIII).
Thalassophis microcephalus Schmidt, 1852: 78,
taf. II (the coast of Java).
Hydrophis leprogaster Dumeril et Bibron in Fis-
cher, 1856: 53 (the type place is unknown).
Hydrophis guentheri Murray 1884: 385 (Karachi,
Pakistan).
Hydrophis rostralis Smith, 1917: 340 (Kuala
Laruth, Perak, Malaysia).
Description. L. 880–930 mm; L./L. cd. 9.8–10.8;
sq. 17–23+30–43; ventr. 220–350; 5–6 maxillary teeth;
the palatinum with 7–8 teeth, the pterygoideum with
9−12 teeth, the dentale with 12–15 teeth; 1 preocular
and 1 postocular plates; 1 large anterior temporal plate;
6 supralabials: 3rd and 4th ones are in contact with an
eye, only 2nd upper labial contacts with prefrontal
plates; 7–8 infralabials, there are no slitlike scales on
last ones; hexangular body scales (except for the first
triens) lay side by side, with 2–3 tubercles or with two-
dented or three-dented keel.
They are grayish above and pale-yellow beneath,
with 40–60 dark crossbands and spots. The head is
olive or yellowish.
Geographical variability. At present two subspe-
cies are admitted to exist: nominative, dwelling on the
majority of the area, and H. gracilis microcephalus
[32], living on the coast of Java, in the Strait of Mal-
acca, and on the coast of Australia and New Guinea.
The latter differs in the bigger number of scales around
the neck (21–23, rarely 19, as against 17–19), around
the body middle (35–43 as against 30–36), and of the
ventrals (250–350 as against 220–287). The existence
of these geographical forms is also corroborated by our
material.
Distribution. From the Persian Gulf and the Ara-
bian Sea in the west to Hong Kong and the coastal
waters of Australia in the east.
Hydrophis (Microcephalophis) cantoris Gunther,
1864
No. 6 2004
390 KHARIN
Hydrophis cantoris Gunther, 1864: 374, pl. XXV,
Fig. 5 (Penang, Malaysia).
Distira gillespiae Boulenger, 1899: 642, pl. (Kara-
chi, Pakistan).
Description. L. 1330–1740 mm; L./L. cd. 10.8–
12.4; sq. 21–25+41–48; ventr. 404–468; 5–6 maxillary
teeth; the palatinum with 6–7 teeth, the pterygoideum
with 6–9 teeth, the dentale with 10–14 teeth; 1 preocu-
lar and 1 postocular; 1 large anterior temporal plate;
6 supralabials: 2nd and 3rd are in touch with eye
(sometimes only 3rd) and with prefrontal plates; 7–8
infralabials; slitlike scales are absent; in dorsal part and
on sides (besides the first quarter) hexangular body
scales lay side by side, with 2–4 tubercles; ventrals with
2–4 tubercles.
Dark brown or gray, with light brown or yellowish
crossbands or lateral spots. The head is black in young
individuals, brown, gray or yellowish green in adults.
Distribution. The coastal waters of Pakistan, India,
Ceylon, and Burma.
Subgenus Mediohydrophis Kharin subgen. nov
Type species: Hydrophis klossi Boulenger, 1912. It
is designated here.
Diagnosis. The maxillare doe not protrude before
palatinum (Fig. 1b). The parietale is always connected
with the praefrontale so that it separates the frontale
from orbital margin and from the postorbitale (Fig. 1e).
Ventrals are entire on the entire interval and are not
divided into two halves in the posterior part of body.
Comparative remarks. Subgenus Mediohydrophis
is intermediate between subgenera Microcephalophis
and Hyrophis s. str. It is indicated by the frontale and
the parietale morphology common for the representa-
tives of subgenera Mediohydrophis and Microcephalo-
phis. The species of subgenus Mediohydrophis have the
same ventral structure and upper jaw not protruding
before the palatinum like the species of subgenus
Hydrophis s. str.
Etymology. The name is formed from two words:
“medio” (lat.) middle, it specifies the intermediate posi-
tion of the subgenera, and “hydrophis” the water snake.
The subgenus includes 2 species.
Hydrophis (Mediohydrophis) klossi Boulenger,
1912.
Hydrophis klossi Boulenger, 1912: 190 (Selengor,
Malaysia).
Description. L. 975–1190 mm; L./L. cd. 8.4–10.8;
sq. 23–27+31–39; ventr. 360–430; 5–6 upper jaw teeth;
palatinum with 6–7 teeth, pterygoideum with
7−10 teeth, dentale with 12–13 teeth; 1 preocular plate
and 1 postocular plate; 1 large anterior temporal plate;
5–6 supralabials: 3rd and 4th are in contact with eye,
2nd is the biggest; 6–7 infralabials; slitlike scales are
absent; body scales are smooth or with small keel; ven-
trals as a rule with 2 tubercles.
RUSSIAN JOURNAL OF MARINE BIOLOGY
They are grayish or greenish above, yellowish
beneath, with 50–75 dark crossbands. The head is usu-
ally with yellow horseshoe marking.
Distribution. The coast of Malaysia, the strait of
Malacca, the gulf of Siam (the coast of Southern Viet-
nam and Campuchia, 7 sp. in the TINRO-Center
museum), Thailand. Thus, the assumption about the
occurrence of this species in Vietnam waters [21] has
completely proven to be true.
Hydrophis (Mediohydrophis) melanosoma Gunther,
1864.
Hydrophis melanosoma Gunther, 1864: 367,
pl. XXV, figure F (the type locality is unknown).
Hydrophis floweri Boulenger, 1898: 106, pl. IX
(Brunei Bay, Kalimantan Island).
Distira wrayi Boulenger, 1900: 307 (Perak, south-
eastern Malaysia).
Description. L. 1000–1390 mm; L./L. cd. 7.4–8.7;
sq. 25–29+37–41; ventr. 266–368; 5–7 upper jaw teeth;
palatinum with 7–8 teeth, pterygoideum with 9–12 teeth,
dentale with 13–14 teeth; 1 preocular plate and 1 pos-
tocular plate; 1 large anterior temporal plate;
6−7 supralabials: 3rd and 4th or 3rd, 4th, and 5th—con-
tact with eye, 2nd, 3rd, and 4th—are the biggest;
7−8 infralabials; after 2 and 3 infralabials slitlike scales
series is present; body scales placed tegularly with
strong keel; ventrals have 2 tubercles.
They are grayish above, with 50–70 wide cross-
bands, the space of which exceeds or is equal to inter-
vals between them. The head and neck are black, with
yellowish narrow, almost oval marking through the bot-
tom of nasals, prefrontals, preoculars, postoculars, and
temporals. Sometimes both branches of this mark
merge on the occiput. Frontals and occipitals can have
5–7 yellow specks.
Distribution. The coast of southeastern Malaysia,
the strait of Malacca, the coast of Kalimantan, Sumatra
Islands, the north of Australia.
Subgenus Hydrophis Latreille in Sonnini et
Latreille, 1802.
Type species: Hydrus fasciatus Schneider, 1799.
Diagnosis. The maxillare is not protruding before
the palatinum (Fig. 1c). The parietale is connected with
the praefrontale incompletely so that the frontale par-
tially enters into orbital margin and contacts with the
postorbitale (figure F). Ventrals are entire, not divided
by the elongate groove in the posterior part of body.
The subgenus includes 7 species.
Hydrophis (Hydrophis) obscurus Daudin, 1803
Hydrophis obscurus Daudin, 1803: 375 (the coast of
Bangladesh, the description is based on: Russell, 1801:
pl VIII). Hydrophis chloris Daudin 1803: 377, pl. XC
(in the same place where is previous, the description is
based on Russell, 1801: pl. VII). Hydrophis subcinctus
Gray, 1842: 63 (India). Hydrophis latifasciata Gunther
1864: 372, pl. XXV, Fig. T (Merguis islands). Hydro-
Vol. 30 No. 6 2004
 REVIEW OF SEA SNAKES
phis coronatus Cunther, 1864: 372, pl. XXV, Fig. M
(the Bay of Bengal).
Description. L. 1055–1090 mm; L./L. cd. 7.8–9.9;
sq. 19–23+25–37; ventr. 298–346; 4–7 upper jaw teeth;
palatinum with 7–8 teeth, pterygoideum with
8−10 teeth, dentale with 14–17 teeth; 1 preocular plate
and 1 postocular plate; 1 large anterior temporal plate;
6–7 supralabials: 3rd and 4th in contact with eye,
1st and 2nd are the smallest; 7–8 infralabials; between
3rd and 4th infralabials slitlike scale is present; body
scales are smooth or with the central two-dented keel,
lay tegularly; ventrals are usually with 2 tubercles.
These sea snakes are grayish or albescent, with
34−60 black or brown crossbands which are much
wider throughout than the grayish intervals between
them. The head is black or dark brown, with a yellowish
horseshoe marking extending from the snout to the
occipitals.
Distribution. The eastern coast of India, the coast
of Ceylon and Burma. All reports for New Guinea [8, 9,
24, 27] apply to species Hydrophis vorisi.
Hydrophis (Hydrophis) vorisi Kharin, 1984.
Hydrophis vorisi Kharin, 1984a: 630, Figs. 1–4 (the
southern coast of New Guinea).
Description. L. 635 mm; L./L. cd. 12.7; sq. 23+29–
35; ventr. 331–345; 6 upper jaw teeth; palatinum with
9 teeth, pterygoideum with 10 teeth, dentale with
14 teeth; 1 preocular plate and 1 postocular plate;
1 large anterior temporal plate: 3rd and 4th ones contact
with eye; 2nd, 3rd, and 4th plates are the biggest;
8 infralabials, 4th and 5th infralabials are separated
from mouth edge by slitlike scales; body scales are
smooth in the first quarter, with the lengthened keel fur-
ther, lie tegularly; ventrals are smooth.
Specimens are brownish, with darker head and
73−74 dark crossbands. The last ones are along the full
extent much wider than the yellowish gray intervals
between them. The head has a yellowish elongate strip
placed in the temporal part and directed through the
corner of the mouth.
Distribution. Hitherto only two specimens of this
species are known from the southern coast of New
Guinea [holotype AMNH 58869, eastern shoal on the
Fly River, and one specimen in the Museum of Com-
parative Zoology (MCZ) NR 141279 – Balimo,
Aramiya River]. It is vicarious species of H. obscurus
from the Australian seas [19].
Hydrophis (Hydrophis) brookii Gunther, 1872.
Hydrophis brookii Gunther, 1872: 597, Fig. 5 (Sara-
vak, Kalimantan Island).
Hydrophis consobrinus Smith, 1917: 341 (the Gulf
of Siam, mouth of the Bankpakong River).
Description. L. 890–920 mm; L./L. cd. 8.0–11.8;
sq. 25–31+37–45; ventr. 328–453; 5 upper jaw teeth;
palatinum with 7–8 teeth, pterygoideum with 14–15
teeth, dentale with 15–16 teeth; 1 preocular plate and
1 postocular plate; 1 large anterior temporal plate;
RUSSIAN JOURNAL OF MARINE BIOLOGY Vol. 30
391
6−7 supralabials: 3rd and 4th ones contact with eye;
2nd, 3rd, and 4th plates are the biggest; 7–8 infralabi-
als; there is a slitlike scale between 3rd and 4th infrala-
bials; body scales are placed almost tegularly, carry the
lengthened keel; ventrals are usually with 2 tubercles.
These snakes are gray above and yellowish white
beneath, with 60–80 crossbands or with large spots (in
the latter case as a rule there are 2 lines of fine spots on
the sides). The crossbands are wider than interspaces
between them on the back and narrower than them
along the sides. The head is dark, with a yellow strip
through a snout and along the sides.
Distribution. The Gulf of Siam, the northern coast
of Java and Kalimantan. All references to the northern
Australian shelf [8, 24, 27] to the species Hydrophis
macdowelli.
Hydrophis (Hydrophis) macdowelli Kharin, 1983
Hydrophis macdowell Kharin, 1983: 1751, Figs. 1–4
(the northern Australian shelf).
Description. L. 706–750 mm; L./L. cd. 7.8–9.2; sq.
23–29+35–44; ventr. 235–290; 6–7 upper jaws teeth;
palatinum with 8 teeth, pterygoideum with 13–15 teeth,
dentale with 15–16 teeth; 1 preocular plate and 1 pos-
tocular plate; 1 large anterior temporal plate;
6−7 supralabials: 3rd and 4th ones are in contact with
eye, they are the biggest; 7–8 infralabials; between 3rd
and 4th infralabials slitlike scale is present; keeled body
scales lay almost tegularly; ventrals are as a rule with
2 little tubercles.
They are cream above and pale yellow beneath, with
36–38 black rhomboid crossbands or spots between
which there are black specks. On the sides there are two
lines of the spots which are the groups of scales with a
black bordering, the head is dark olive.
It differs significantly from all the species of the
genus (including the close H. brookii) by the palatinum
structure without prominence for the maxillare.
Distribution. The shelf of Northern Australia, the
southern coast of New Guinea. It is the geographical
vicarious species of H. brookii in the Australian waters
[18].
Hydrophis (Hydrophis) fasciatus [31].
Hydrus fasciatus Schneider, 1799: 240 (the coast of
East India, part.).
Hydrophis laticauda Latreille in Soninni, Latreille,
1802: 195 (the coast of China, part.).
Aturia lindsayi Gray, 1842: 61 (the coast of China).
Hydrophis leptodira Boulenger, 1896: 285 (the
mouth of the Ganges).
Distira rombifer Boulenger, 1900: 306 (Perak,
Malaysia).
Description. L. 815–1010 mm; L./L. cd. 10.1–10.8;
sq. 27–33+47–58; ventr. 410–514 (usually no more
than 460); 5–6 upper jaw teeth; palatinum with 7–8
teeth, pterygoideum with 8–10 teeth, dentale with
13−15 teeth; 1 preocular plate and 1, rarely 2, postocu-
No. 6 2004
392 KHARIN
lars; 1 large anterior temporal plate; 5–7 supralabials:
3rd and 4th ones contact with eye, two first plates are
very small; 6–7 infralabials; between 3rd and
4th infralabials slitlike scale is present; body scales lay
almost tegularly, they are hexangular in the middle and
posterior parts, with central tubercle or short keel; ven-
trals as a rule are smooth.
Head, neck, and usually anterior body part are black
or dark olive with pale yellow oval specks on each side
of the body. The posterior part of these specimens is
grayish above and albescent beneath, with dark rhom-
boidal spots or crossbands on the back (especially in
young individuals).
Distribution. The coastal waters of Pakistan, India,
Ceylon, Burma, Singapore, the Strait of Malacca, the
coastal waters of China, Java. The specimen from the
northern Australian shelf (the Australian museum -AM
R 14022) has 484 ventrals [8] and should be related to
this species.
Hydrophis (Hydrophis) atriceps Gunther, 1864.
Hydrophis atriceps Gunther, 1864: 371, pl. XXV,
Fig. A (Thailand).
Hydrophis alcoki Wall, 1906: 288, Fig. (Puri,
Orissa, India).
Disteira cincinatii Van Denburgh, Thompson, 1908:
41, pl. X (Philippines).
Description. L. 820–1015 mm; L./L. cd. 10.2–0.7;
sq. 25–31 (more often 27–29) +39–49 (more often 43–
45); ventr. 323–453 (as a rule 366 and fewer);
5−6 upper jaw teeth; palatinum with 7–8 teeth, ptery-
goideum with 8–13 teeth, dentale with 13–15 teeth;
1 preocular plate and 1–2 postoculars; 1 large anterior
temporal plate; 6–7 supralabials: 3rd and 4th ones con-
tact with eye, first one or two are very small;
7−8 infralabials; between 3rd and 4th infralabials there
is a slitlike scale; body scales are placed almost tegu-
larly, they are hexangular in the middle and posterior
parts, with a central tubercle or short keel; ventrals are
usually smooth.
Body coloration is the same as at the previous spe-
cies. Individuals with well-pronounced crossbands
occur more often than in the preceding species,
50−75 crossbands. On the head behind the nostrils and
eye or behind only the eye a yellow spot can be present.
Distribution. The coast of India (holotype
H. alcocki), Burma, the Gulf of Siam, the coast of the
Java and the Aru Islands, Philippines, the coast of
China (ZMB 9697), and Northern Australia (AM
6715).
Comparative remarks. Up until recently it was
considered a subspecies of H. fasciatus. It is however
seen from the distribution to dwell together with H. fas-
ciatus on the most part of its areal, what contradicts
subspecies criterion [25]. Additionally, it differs con-
siderably from H. fasciatus by the lesser quantity of
ventrals [370.3 (n = 14—our material and n = 32—
Smith’s data [35]) as against 464.2 (n = 17—our mate-
RUSSIAN JOURNAL OF MARINE BIOLOGY
rial and n = 29—Smith’s data [35])] and also of the
scales around the neck (27.5 as against 30.7) and body
middle (44.2 as against 53.1). On the base of this we
consider H. atriceps as a separate species that fully cor-
respond with the annotated catalogue of poisonous
snakes [13].
Hydrophis (Hydrophis) parviceps Smith, 1935
The material is absent. The description is based on
two specimens reported by Smith (Smith, 1935: 5,
Fig. 2; 1943: 465, Kohinchina coast, 9°59′ N and
107°23.5′ E) and Taylor (Taylor, 1963: 432, South
China Sea, 10°17′ N and 107°47′ E). Holotype is stored
in the Zoological Museum of Copenhagen, Denmark,
ZMUC R 66182 [42].
Description. L. 765–890 mm; L./L. cd. 8.2–8.3; sq.
19–21+31–33; ventr. 340–348; 6 upper jaw teeth; the
palatinum with 7 teeth; the number of teeth on the
pterygoideum and the dentale is unknown; 1 preocular
plate and 1 postocular plate; 1 large anterior temporal
plate; 5 supralabials: 3rd and 4th ones contact with eye,
2nd plate is the biggest; 7 infralabials; slitlike scales are
absent on them; hexangular body scales lay almost teg-
ularly, smooth in anterior part and keeled in middle and
posterior parts; ventrals are with two big keels, often
with 1 or more tubercles.
These sea snakes are grayish olive, with 69 black
crossbands which are wider than intervals between
them on the back and narrower than them on the sides.
The head is black, with a narrow yellowish marking
through preoculars and prefrontals. The body is albes-
cent beneath.
Distribution. It is known from two specimens from
the South China Sea.
Remark. In spite of lacking data on the osteology
we refer this species to the subgenus Hydrophis s. str
conventionally after McDowell [24]. H. parviceps is
seen from the description to be very close to H. atriceps
on morphology, which has been emphasized both by
the author of this species [36, 37] and by Taylor [40]
who examined the second specimen.
ACKNOWLEDGMENTS
The author is sincerely grateful to V.P. Shuntov
(TINRO) for the constant help during the work, and
also to I.S. Darevskii (ZIN), E.N. Arnold and
K. McCarthy (BM), R.G. Tsveifel (AMNH), R. Gunter
(ZMB), and A.R. Rasmussen (ZMUC) for kindly
loaned material.
REFERENCES
1. Boie, F. von., Bemerkungen über Merrem’s Versuch
eines Systems der Amphibien, Ibis, 1827, Bd. 20,
S. 1−12, pp. 508–566.
2. Boulenger, G.A., The fauna of British India, including
Ceylon and Burma, Reptilia and Batrachia, London:
Taylor and Francis, 1890.
Vol. 30 No. 6 2004
 REVIEW OF SEA SNAKES
3. Boulenger, G.A., Catalogue of the Snakes in the British
Museum. 3, Containing the Colubridae (Opistoglyphae
and Proteroglyphae), Amblycephalidae, and Viperidae, 22.
London: Taylor and Francis, 1896.
4. Boulenger, G.A., Description of a New Sea Snake from
Borneo, Proc. Zool. Soc. Lond., 1898, pp. 106–107.
5. Boulenger, G.A., A New Sea Snake of the Genus Distira 23.
from Kurrachee, J. Bombay Nat. Hist., 1899, vol. 12,
p. 642.
6. Boulenger, G.A., Descriptions of New Reptiles from
Perak, Malay Peninsula, Ann. Mag. Nat. Hist., 1900,
Ser. 7, Vol. 5, pp. 306–308.
7. Boulenger, G.A., A Vertebrate Fauna of the Malay Pen-
insula from the Isthmus of Kra to Singapore Including
Adjacent Islands, Reptilia and Batrachia, London, 24.
1912.
8. Cogger, H.G., Sea Snakes of Australia and New Guinea,
The Biology of Sea Snakes Baltimore, London: Univ. 25.
Park. Press., 1975a, pp. 59–139.
9. Cogger, H.G., Reptiles and Amphibians of Australia, 26.
Sydney: A. and W. Reed, 1975b.
10. Daudin, F.M., Histoire Naturelle Generale et Particu-
liere des Reptiles 7, Paris, 1803. 27.
11. Dumeril, A.M.C., Bibron, G., and Dumeril, A., Erpetol-
ogie Generale ou Histoire Naturelle Complete des Rep-
tiles, 7–10, Paris, 1854, pp. 781–1536. 28.
12. Fischer, J.G., Die Familie der Seeschlangen, System-
atisch Beschrieben, Abh. Naturw. Ver., Hamburg, 1856,
29.
Bd. 3, pp. 1–78.
13. Golay, P., Smith, H.M., Broadley, D.G., et al.,
Endoglyphs and Other Major Venomous Snakes of the
World, A Checklist, Geneva: S.A. Azemiops Herpetolog- 30.
ical Data Center, 1993.
14. Gray, J.E., Monographic Synopsis of the Water Snakes
of the Family Hydridae, Zool. Misc., 1842, vol. 4,
pp. 59–68.
15. Günther, A., The Reptiles of British India, London: Rob- 31.
ert Hardwick Publ., 1864.
16. Günther, A., On the Reptiles and Amphibians of Borneo,
Proc. Zool. Soc. Lond., 1872, pp. 586–600.
17. Kharin, V.E., The Review of Sea Snakes of the Genus 32.
Aipysurus (Serpentes, Hydrophiidae), Zool. Zhurn.,
1981, vol. 60, issue 2, pp. 257–264.
18. Kharin, V.E., The New Species of Sea Snakes of the 33.
Genus Hydrophis sensu lato (Serpentes, Hydrophiidae)
from the Northern Australian shelf, Zool. Zhurn., 1983,
vol. 62, issue 11, pp. 1751–1754.
19. Kharin, V.E., The Sea Snakes of the Genus Hydrophis 34.
sensu lato (Serpentes, Hydrophiidae). 2. To the Question
About the Taxonomical Position of H. obscurus from
New Guinea, Zool. Zhurn., 1984a, vol. 63, issue 4, 35.
pp. 630–632.
20. Kharin, V.E., The Review of Sea Snakes of the Hydro- 36.
phis Group sensu lato (Serpentes, Hydrophiidae). 3. The
Genus Leioselasma, Zool. Zhurn., 1984b, vol. 63, issue 10, 37.
pp. 1535–1546.
21. Kharin, V.E., About Three Species of the Sea Snakes
First Occurred in Vietnam Waters, with the Remark on
RUSSIAN JOURNAL OF MARINE BIOLOGY Vol. 30 No. 6
393
Rare Form Praescutata viperina, Biol. Morya, 1984,
no. 2, pp. 26–30.
Kharin, V.E., About the Taxonomical Position of Sea
Snake Hydrophis caerulescens (Shaw, 1802) (Serpentes:
Hydrophiidae), Biol. Morya, 2004, vol. 30, no. 3,
pp. 227–229.
Lesson, R.P., Reptiles, Voyage aux Indes Orientales par
le Nord de l’Europe, les Provinces du Caucase, la Geor-
gie, l’Armenie et la Perse, suivi de détails
topographiques ttatistiques et autres sur le Pégou, les
Iles de Java, de Maurice et de Bourbon, sur le Cap-de-
Bonne-Esperance et Sainte-Héléne, pendant les années
1825–1829, Zoologie, Paris: Arthur Bertrand, 1834,
pp. 291–336.
McDowell, S.B., The Genera of Sea Snakes of the
Hydrophis Group (Serpentes, Elapidae), Trans. Zool.
Soc., London, 1972, vol. 32, pt. 3, pp. 195–247.
Mayr, E., Printsipy zoologicheskoi sistematiki (Princi-
ples of Zoological Systematics), Moscow: Mir,1971.
Mengden, G.A., The Taxonomy of Australian Elapid
Snakes. A Review, Rec. Austral. Mus., 1983, vol. 35,
pt. 5–6, pp. 195–223.
Minton, Sh.A., Geographic Distribution of Sea Snakes,
The biology of sea snakes, Baltimore; London: Univ.
Park. Press., 1975, pp. 21–30.
Murray, J.A., The Vertebrate Zoology of Sind, Bombay,
1884.
Rasmussen, A.R., Systematics of Sea Snakes: a Critical
Review, Venomous Snakes-Ecology, Evolution and
Snake Bite, Oxford: Clarendon Press., 1997, pp. 15–30.
Russell, P., A Continuation of an Account of Indian
Serpents: Containing Descriptions and Figures from
Specimens and Drawings, Transmitted from Various
Parts of India to the Hon. Court of Directors of the
East India Company, London: W. Bulmer and Co.
1801, vol. 2.
Schneider, I.G., Historiae amphibiorum naturalis et liter-
ariae falciculus primus continens Ranes, Calamitas,
Bufones, Salamandras et Hydros genera et species
descriptos notisque suis distinctos, Jennae, 1799.
Schmidt P., Beitrage zur Fernern Kentniss der Meer-
schlangen, Abh. Naturw. Ver., Hamburg, 1852, Bd. 2,
pp. 72–86.
Shaw, G., General Zoology or Systematic Natural His-
tory by George Shaw, M.D.F.R.S., with Plates from the
first Authorities and Most Select Specimens. Amphibia,
London, 1802, vol. 3, pt. 2.
Smith, M.A., Preliminary Diagnoses of Four New Sea
Snakes, J. Nat. Hist. Siam., 1917, vol. 2, pp. 340–342.
Smith, M.A., Monograph of the Sea Snakes (Hydrophi-
idae), London, 1926.
Smith, M.A., The Sea Snakes (Hydrophiidae), Dana
Rept., 1935, no. 8, pp. 1–6.
Smith, M.A., The Fauna of British India, Ceylon and
Burma, Including the Whole of the Indo-Chinese Subre-
gion, Reptilia and Amphibia, London: Taylor and Fran-
cis 1943, vol. 3: Serpentes.
2004
394 KHARIN
38. Sonnini, Ch.S., Latreille, P.A., Histoire Naturelle des
Reptiles, avec figures dessinees d’apres nature, Paris:
Deterville, 1802, vol. 4.
39. Stejneger, L., Herpetology of Japan and Adjacent Terri-
tory, U.S. Nat. Mus. Bull., 1907, vol. 58, pp. 1–577.
40. Taylor, E.H., New and Rare Oriental Serpents, Copeia,
1963, no. 2, pp. 429–433.
41. Van Denburgh, J., Thompson, I.C., Description of a New
Species of Sea Snake from the Philippine Islands, with a
Note on the Palatine Teeth in the Proteroglypha, Proc.
Calif. Acad. Sci., 1908, ser. 4, vol. 3, pp. 41–48.
42. Voris, H.K., The Role of Sea Snakes (Hydrophiidae) in
Trophic Structure of Coast Ocean Communities, J. Mar.
Biol. Assoc., India, 1972, vol. 14, pt. 2, pp. 429–442.
RUSSIAN JOURNAL OF MARINE BIOLOGY
43. Voris, H.K., A phylogeny of the Sea Snakes (Hydrophi-
idae), Field Zool., 1977, vol. 70, pt. 4, pp. 79–169.
44. Wall, F., A Descriptive List of Sea Snakes (Hydrophii-
nae) in the Indian Museum, Calcutta, Mem. Asiat. Soc.,
Bengal, 1906, vol. 1, no. 14, pp. 277–299.
45. Wall, F., A Monograph of the Sea Snakes (Hydrophii-
nae), Mem. Asiat. Soc. Bengal, 1909, vol. 2, no. 8,
pp. 169–251.
46. Wall, F., Ophidia taprobanica or the Snakes of Ceylon,
Colombo: H.R. Cottle, 1921.
47. Welch, K.G.R., Snakes of the World: A checklist, 1994,
vol. 1: Venomous snakes.
Vol. 30 No. 6 2004