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THE RAFFLES BULLETIN OF ZOOLOGY 2000 Supplement No.

8: 459-493
National University of Singapore

ECHINODERM FAUNA OF THE SOUTH CHINA SEA:


AN INVENTORY AND ANALYSIS OF
DISTRIBUTION PATTERNS
David J.W. Lane
Department of Biological Sciences, National University of Singapore, Lower Kent Ridge Road, Singapore
119260, Republic of Singapore. E-mail: dbslaned@nus.edu.sg

Loisette M. Marsh
Department of Aquatic Zoology, Museum of Natural Science, Western Australian Museum, Francis Street,
Perth, Western Australia 6000, Australia

Didier VandenSpiegel
Musee Royal de I'Afrique Centrale, Invertebres non insectes, B-3080 Tervuren, Belgium

Frank W.E. Rowe


Goldbrook Boarding Kennels, Nuttery Vale, Cross Street, Hoxne, Suffolk, IP21 5BB,

u.K.

ABSTRACT. - A comprehensive review and analysis of the literature on echinoderm

records for the South China Sea (SCS) indicates close to a thousand (982) species in
total (113 crinoids, 227 asteroids, 272 ophiuroids, 167 echinoids and 203 holothuroids).
All known SCS species and their distributions are tabulated herein. A total of 178
echinoderms have their type locality in the South China Sea, with 63% of these (12%
of the echinofauna overall) currently considered endemic. One possible reason for the
prominence of endemics is that the South China Sea became relatively land-locked,
repeatedly, during low sea level stands. Large areas of the South China Sea remain
relatively unexplored biologically and it is likely that additional records and new taxa
await discovery.

KEY WORDS. - Echinodermata, South China Sea, Indo-Pacific, biodiversity,


biogeography

INTRODUCTION
The fauna of the South China Sea has never been comprehensively studied and most
expeditions have merely touched the fringes. The first major expedition to collect in the
South China Sea was the renowned world-wide, deep-sea exploring expedition of HMS
"Challenger" (1873-76). "Challenger" occupied many stations through the Philippines but
only two in the South China Sea, one off the west coast of Luzon and one near Hong Kong.

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Echinoderms from the voyage were described by Sladen, 1889 (asteroids), Lyman, 1882
(ophiuroids), Theel, 1882, 1886 (holothurians), Carpenter 1884, 1888 (crinoids) and A.
Agassiz, 1881 (echinoids). Bedford (1900) listed echinoderms from Singapore and peninsular
Malaysia and A.H. Clark (1934) described the crinoids in the Raffles Museum. Bell (1894)
described the echinoderms collected from Macclesfield Bank by HMS "Penguin" and HMS
"Egeria". By far the most significant contribution to the marine zoology of the area was
made by the United States Fisheries steamer "Albatross" (1907-1910) which dredged through
the Philippines and adjacent areas but again only touched the fringes of the South China
Sea, from Balabac Strait, the north west coast of Palawan, the deep Palawan passage and
a number of stations north west of Mindoro and off the west coast of Luzon. The echinoderms
were described by Fisher, 1919 (asteroids), Koehler, 1922 (ophiuroids), A.H. Clark, 1911
(crinoids) and Mortensen, 1927, 1940b, 1948b (echinoids). Domantay (1933, 1934, 1936,
1962, 1972), Domantay & Domantay (1967) and Domantay & Roxas (1938) contributed to

20

t---+------+-+----cA=\----+-----+----.:..j~----J

100

20

120

Fig. 1. Map of the South China Sea. SCS boundaries and demarcations between the seven zones (17) are indicated with dashed lines. MB = Macclesfield Bank.

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knowledge of the echinoderms of the coasts of Mindoro and Luzon. More recently Liao has
made extensive studies of the echinoderm fauna of China from both deep and shallow water
including the Paracel (Xisha) Islands, culminating in Liao & Clark (1995) and Liao (1997).
Chinese scientists have made a substantial contribution to knowledge of the Spratly (Nansha)
islands and neighbouring waters, including deep water species. Their major report (Guozhen,
1989) lists 162 species, with a further 35 added in a supplementary list (Guozhen, 1991).
Taiwan echinoderms have been studied principally by Chao & Chang (1989a, b, 1990) and
Applegate (1984). Vietnam echinoderms are listed by Dao (1991a, b, c, 1994) and Cherbonnier
(1960, 1961). The French MUSORSTOM Expeditions 1 and 2 (off the west coast of the
Philippines) sampled echinoderms from the shelf and slope to 970 metres (Guille, 1981;
Bourseau & Roux, 1989).
Comprehensive works by Mortensen (Monographs of the Echinoidea - 1928, 1935, 1940a,
1943, 1948a, 1950, 1951), A.H. Clark (Monograph of the existing crinoids 1931, 1941, 1947,
1950; AH. Clark & AM. Clark, 1967) and AM. Clark & Rowe (1971) have brought together
diverse records and have largely stabilised the taxonomy. Rowe & Gates (1995), who list
all species known from Australia, have updated the taxonomy, including many species also
found in the South China Sea.
The South China Sea (SCS) is largely enclosed by major land masses and island chains,
with more open water in the Luzon Strait to the north and between Kalimantan and Sumatra
to the south. For the purposes of this report the northern Luzon Strait boundary is designated,
arbitrarily, from Escarpada point, N. Luzon, to S.E. Taiwan, including the Babuyan and
Batan Islands. The southern boundary, at about 3 south of the equator, includes the islands
of P. Bangka and P. Belitung between Sumatra and Kalimantan. Similarly, islands within
the Balabac Strait, the Linapacan / Mindoro Straits and Verde Island Passage are considered
included by the eastern Philippine boundary of the SCS. The islands at the southern end
of the Straits of Malacca are also included (Fig. 1). For the purposes of the distribution
tables in this report the area is subdivided into seven zones, 1 to 4 being coastal, 5 and 6
more central, and area 7 encompassing shallow and deep waters of the Nansha (Spratly)
islands (Fig. 1). The echinoderm inventory includes all known records published in the
literature. Also included are data from unpublished reports of the second author and from
field observations / collections of the first author (the latter indicated by L in distribution
table). Depth ranges for non-endemic taxa include records from outside as well as within
the South China Sea. An asterisk instead of a plus sign in column 3 of the table indicates
either an SCS taxon recorded for the adjacent central Philippine Archipelago/Sulu Sea (but
not in area 3), or an unspecified locality in the Philippines. This is considered useful since
past Philippines expeditions have largely concentrated on the central Philippines-Sulu region;
representation in area 3 might be expected for species which range from central archipelagic
waters to other SCS zones. South China Sea type species and their type zones, where known,
are highlighted in bold in the distribution tables. Although the present lists are as
comprehensive as possible, some records may have been overlooked, and further taxonomic
work, needed for many taxa, may reveal synonymies or result in the recognition of additional
species.

ZOOGEOGRAPHIC ANALYSIS
The present lists comprise 982 species of echinoderms (113 crinoids, 227 asteroids, 272
ophiuroids, 167 echinoids and 203 holothurians). For comparison Rowe & Gates (1995) list

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1154 species for Australia waters, with about two thirds of these comprising non-endemic
and endemic, tropical species (Rowe, 1985). To our knowledge no compilation has been
made of the total echinoderm fauna (i.e. at all depths) of the species rich, core Indo-Malay
Archipelago area, i.e. the remainder of the Philippines, including the Sulu Sea, the Indonesian
Archipelago, New Guinea and islands to the north of New Guinea. This area probably has
a richer fauna than the South China Sea but no comparisons, other than of shallow-water
30m) species (c.f. Clark & Rowe, 1971), can be made at present; however it is known that
the Sulu Sea, because of its historical isolation during successive Pleistocene glaciations,
has a high level of faunal endemicity at species level.
As results from the MUSORSTOM expeditions show, viz. their recently described species
from the eastern part of the South China Sea, there are probably still many species to be
discovered in other deep water areas. In general, many deep water species are known only
from a single collection, sometimes from a single specimen; these are considered to be
endemic until shown to have a wider distribution. On the other hand some deep water species,
living at low temperatures, have a very wide distribution through the Indo-Pacific and
sometimes the Atlantic, at temperate as well as tropical latitudes.
The species distributions fall quite neatly into the following zoogeographical categories: (1)
endemic to the South China Sea; (2) confined to all or part of Southern and South East Asia
from India/Sri Lanka coastal waters to Japan, south to Indonesia and northern Australia; (3)
those found in this area but extending either into the Indian or Pacific oceanic domains,
(restricted IWP in Table 1) and (4) those found widely distributed in the Indo-West Pacific
(IWP), sometimes from East Africa to Hawaii. A few cosmopolitan species are included
in the IWP group (Table 1).

Table 1. Distribution patterns of echinoderms from the South China Sea.


Classes

Total species

Crinoidea

113

Asteroidea

Endemic
SCS

S. Asia

Australia

Restricted
IWP

(7%)

67 (63%)

24 (22%)

227

54 (24%)

80 (36%)

55 (24%)

IWP
9

(8%)

36 (16%)

Ophiuroidea

272

13

(5%)

132 (49%)

68 (26%)

54 (20%)

Echinoidea

167

(5%)

68 (45%)

36 (24%)

42 (28%)

Holothuroidea

203

31 (16%)

51 (27%)

43 (23%)

64 (34%)

Totals

982

112 (12%)

398 (42%)

226 (24%)

205 (22%)

A total of 178 echinoderms (18% of the fauna) have their type locality in the South China
Sea or on the coasts bordering it: 14 crinoids, 61 asteroids, 41 ophiuroids, 19 echinoids and
43 holothurians. About 63% of these are currently considered endemic species (see Table
1) based on the present state of knowledge of their geographic distribution and, for some,
their taxonomy.
It should be noted that distribution data was not available for all species, hence the figures

do not add up to the total number of species. Percentages were calculated from the number
of species for which data was available; no distribution data was available for 6 crinoids,
5 ophiuroids, 15 echinoids and 14 holothurians (2 asteroids plus 2 additional holothurians,
identified to genus only, were also excluded from the analysis). Nevertheless it is believed

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that the data, overall, do give a fairly accurate picture of the patterns of distribution.
The rate of endemicity in the South China Sea is quite high, comprising 12% of the
echinofauna overall and 24% and 16% for asteroids and holothuroids respectively. Endemic
South China Sea echinoderms account for 63% of those with type locality in the region. One
possible reason for the prominence of endemics is that the South China Sea became relatively
land-locked periodically during glacial low sea level stands. The highest proportion of species
(42%) is widely distributed to parts of southern and eastern Asia, often including Northern
Australia. Many fewer species (24 and 22% respectively) are more widely distributed to
the Indian or Pacific oceans (restricted IWP), or to both (IWP). For comparison, re-working
the data in Rowe (1985) and Rowe & Gates (1995) for the Australian tropical fauna gives
the following approximate distribution pattern of species: Endemics - 27%; shared South
East Asian ('East Indian' - includes N. Australian component) - 15%; 'East Indian' plus
either Pacific or Indian Ocean (equivalent to restricted IWP) - 16%; Indo-Pacific (= IWP)
- 21 % (based on a tropical element of 0.67 x 1154 species).
One area in the South China Sea which is conspicuously lacking in data for echinoderms
(and probably other groups) is the Nansha (Spratly) islands reefal system. The present
echinoderm count of 205 species for this area is relatively low, given the complexity and
size of this shallow to deep tropical area. Further sampling in these waters will undoubtedly
reveal additional taxa, some of which may boost the endemic component of the South China
Sea. This is also likely to be the case for the deep echinoderm fauna generally.
It is not practical at present to comment on rare species since rarity may be only apparent
because of lack of collecting, however records from a single locality in the species lists
suggest possibly rare species.

Need for further expeditions / surveys


As can be seen from the historical account, most of the major expeditions visiting the South
China Sea were undertaken during latter part of the 19th and early part of the 20th century.
The survey coverage of this area was only partial and large areas of the South China Sea
remain almost completely unknown biologically. In recent years Thai biologists have sampled
echinoderms from the Gulf of Thailand but results are as yet unpublished; only echinoids
are well represented in the literature for this area (Area 1). The small domains of Singapore
and Hong Kong waters have been repeatedly sampled and their echinofauna is relatively
well known, yet the extensive area of continental shelf (part of the Sunda shelf) between
the Thailand-Malaysia peninsula and Borneo is, apart from the east coast islands of
peninsular Malaysia and the Anambas islands, apparently unsampled, as is the deep water
area (South China Basin) between Macclesfield Bank and the large area of shoals to the
north of East Malaysia and Palawan. The coastal fauna of Sarawak and Brunei is little known,
while that of Sabah is slightly better known. While the "Albatross" sampled deep water off
Palawan they collected from few inshore stations on that island. There is also much to be
known of the shoals which include the Spratly Islands (Nansha), Itu Aba (Taiping Island)
and adjacent areas.
Thus it is apparent that much sampling of offshore reef systems (e.g. Nansha islands), nonreefal coasts and deep waters (below 200m) remains to be done before the fauna of the
South China Sea is as well known as that of the Philippines or Indonesia. As a first initiative,
an international expedition to the Nansha islands would be expected to be particularly fruitful
in terms of biodiversity / biogeographic discoveries.

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ACKNOWLEDGEMENTS

The authors thank D.L. Pawson for comments on an earlier version of this manuscript, and
Cai Yixiong and Benito Tan for assistance translating Chinese publications.

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468

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from the Red Sea. Beaufortia, 48(3): 27-46.

469

Lane et al.: Echinoderm fauna of the South China Sea

ECHINODERM SPECIES LISTS AND DISTRIBUTIONS


Explanation of symbols: Columns 1-7 correspond to zones in Fig. 1; names of SCS type
species are bolded; + =presence of species (bolded for type zones); * = an SCS/Philippines
taxon with status unknown for area 3; L = field observations/collections of the first author.
Distribution

Taxa
2

Depth range m.

+
+
+

+
+
+
+
+

ASTEROIDEA
Order PAXILLOSIDA

Luidiidae
Luidia aspera Sladen, 1889
Luidia avicularia Fisher, 1913
Luidia hardwicki (Gray, 1840)
Luidia longispina Sladen , 1889
Luidia maculata MUller & Troschel, 1842
Luidia magnifica Fisher, 1906
Luidia orientalis Fisher, 1913
Luidia penangensis de Loriol, 1891
Luidia quinaria von Martens, 1865
Luidia savignyi (Audouin, 1826)
Astropectinidae
Astromesites compactus Fisher, 1913
Astropecten acanthifer Sladen , 1883
Astropecten andersoni Sladen, 1888
Astropecten bengalensis DOderlein, 1917
Astropecten debilis Koehler, 1910
Astropecten carcharicus fonnosanus
Diiderlein, 1917
Astropecten eucnemis Fisher, 1919
Astropecten fasciatus Diiderlein, 1926
Astropecten indicus DOderlein, 1888
Astropecten kagoshimensis de Loriol, 1899
Astropecten luzonicus Fisher, 1913
Astropecten malayanus Diiderlein, 1917
Astropecten mindanensis DOderlein, 1917
Astropecten monacanthus Sladen, 1883
Astropecten novaeguineae Diiderlein, 1917
Astropecten polyacanthus phragmorus
Fisher, 1913
Astropecten polyacanthus polyacanthus
MUller & Troschel, 1842
Astropecten pusillus Sluiter, 1889
Astropecten sarasinorum Diiderlein, 1917
Astropecten scoparius, MUller & Troschel, 1842
Astropecten tamilicus Diiderlein, 1888
Astropecten umbrinus Grube, 1866
Astropeeten vappa vappa MUller & Troschel, 1843
Astropecten velitaris von Martens, 1865
Craspidaster hesperus (MUller & Troschel, 1840)
Ctenophoraster diploctenius Fisher, 1913
Ctenopleura astropectinides Fisher, 1913

+
+
+
+

+
+
+

...
+
+

...

314-384
27-256
6-18
9-122
676-765

+
+
+
+

4-274
9-275
6-220
10-148
0-150
18-133
200-380
4-20
0-218
0-50

+
+

...

..
+

+
+

0-250

+
+
+
+
+

+
+

+
+

+
+
+

+
+
+
+

+
+

"'+"

...

+
+
+

181-196
39-119
324-430
216
0-13
0-112
0-88
0-85

0-144
15-22
0-20
14-64
6-15
0-80
5-183
2-195
119-215
90-490

possibly a synonym of Astropecten vappa vappa MUller & Troschel, 1842 (see Liao & Clark, 1995: 77)
although A. carcharicus from type locality (Shark Bay, Western Australia) is apparently distinct from A.
vappa from the same locality (L.M.M.).

470

THE RAFFLES BULLETIN OF ZOOLOGY 2000 Supplement No. 8


Taxa

Distribution
2

Ctenopleura ludwigi (de Loriol, 1899)


Ctenopleura sinica (Doderlein, 1917)
Dipsacaster imperialis Fisher, 1917
Dipsacaster pretiosus (Doderlein, 1902)
Koremaster evaulus evaulus (Fisher, 1913)
Koremaster evaulus spiculatus Fisher, 1919
Persephonaster cingulatus multicinctus
Fisher, 1913
Persephonaster habrogenys Fisher, 1913
Persephonaster tenuis Fisher, 1913
Proserpinaster anchistus (Fisher, 1913)
Proserpinaster euryactis (Fisher, 1913)
Proserpinaster euryactis brevispinus
(Fisher, 1919)
Proserpinaster luzonicus (Fisher, 1913)
Tethyaster aulophora (Fisher, 1911)

Depth range m.
6

+
+

...

+
+

...

...
...
...

685-1020
588-622
134-595
336-432
265-362
314-472

..

...
...

Porcellanasteridae
Abyssaster planus (Sladen, 1883)
Eremicaster crassus (Sladen, 1883)
Thoracaster cylindratus Sladen, 1883
Goniopectinidae
Ctenodiscus orientalis Fisher, 1913
Goniopecten asiaticus Fisher, 1913

10-36
77-134
40-866
20-200
750-1525
1363

420
156-232

+
+
+

3500
1570-6330
2450-5990
605-1360
170-200

+
+

Order NOTOMYOTIDA
Benthopectinidae

..

Cheiraster capillatus Jangoux, 1981


Cheiraster gazellae Studer, 1883
Cheiraster inops Fisher, 1906
Cheiraster niasicus Ludwig, 1910
Cheiraster pilosus (Alcock, 1893)
Cheiraster trullipes (Sladen, 1889)
Nearchaster musorstomi Aziz & Jangoux, 1985
Pectinaster mimicus palawanensis Fisher, 1919

+
+
+
+

.....
..

170-200
170-469
430-1250
170-1080
150-1310
1920
970
1281-2013

Order VALVATIDA
Sphaerasteridae
Podosphaeraster polyplax
A.M. Clark & Wright, 1962
Archasteridae
Archaster angulatus Muller & Troschel, 1842
Archaster typicus Muller & Troschel, 1840

Goniasteridae
Anthenoides cristatus (Sladen, 1889)
Anthenoides granulosus Fisher, 1913
Anthenoides laevigatus Liao & A.M. Clark, 1989
Anthenoides lithosorus Fisher, 1913
Anthenoides rugulosus Fisher, 1913
Anthenoides tenuis Liao & A.M. Clark, 1989
Astroceramus lionotus Fisher, 1913
Astrothauma euphylacteum Fisher, 1913
Calliaster corynetes Fisher, 1913
Calliaster childreni Gray, 1840
Calliaster quadrispinus Liao, 1989

+
+

+
+

...

72-125

+
+

0-90
0-91

.....

+
+

...

..

471

..

90-511
485-526
107-170
380
194-472
180-270
379-407
180-296
196-407
121-192
88

Lane et al.: Echinoderm fauna of the South China Sea


Distribution

Taxa
2

Calliderma emma Gray, 1847


Ceramaster smithi Fisher, 1913
lconaster elegans Jangoux, 1981
[conaster longimanus (Moebius, 1859)
Lithosoma actinometra Fisher, 1911
Milteliphaster spinosus H.L. Clark, 1916
Nymphaster arthrocnemis Fisher, 1913
Nymphaster meseres Fisher, 1913
Nymphaster mucronatus Fisher, 1913
Ogmaster capella (Muller & Troschel, 1842)
Paragonaster ctenipes ctenipes Sladen, 1889
Paragonaster ctenipes hypacanthus Fisher, 1913
Paragonaster stenostichus Fisher, 1913
Peltaster cycloplax Fisher, 1913
Perissogonaster insignis Fisher, 1913
Pontioceramus grandis Fisher, 1911
Pseudarchaster jordani Fisher, 1906
Pseudarchaster mozaicus Wood-Mason &
Alcock, 1891
Pseudoceramaster regularis Jangoux, 1981
Rosaster bipunctus (Sladen, 1889)
Rosaster confinis (Koehler, 1910)
Rosaster mimicus Fisher, 1913
Rosaster symbolicus (Sladen, 1889)
Sphaeriodiscus scotocryptus Fisher, 1913
Stellaster convexus Jangoux, 1981
Stellaster childreni Gray, 1840
Oreasteridae
Anthenea aspera DOderlein, 1915
Anthenea chinensis Gray, 1840
Anthenea difficilis Liao, 1995
Anthenea flavescens (Gray, 1840)
Anthenea grayi Perrier, 1875
Anthenea regalis Koeh1er, 19102
Anthenea viguieri Doderlein, 1915 2
Choriaster granulatus Lutken, 1869
Culcita novaeguineae Muller & Trosche1, 1842
Goniodiscaster forficulatus (Perrier, 1875j3
Goniodiscaster granuliferus (Gray, 1847)
Goniodiscaster pleyadella (Lamarck, 1816)
Goniodiscaster rugosus (Perrier, 1875)
Goniodiscaster scaber (Moebius, 1859)
Gymnanthenea globigera (Doderlein, 1915)
Gymnanthenea laevis H.L. C1ark, 1938
Halityle regularis Fisher, 1913
Monachaster sanderi (Meissner, 1892)
Pentaceraster affinis (Muller & Trosche1, 1842)
Pentaceraster alveolatus (Perrier, 1875)
Pentaceraster chinensis (Gray, 1840)
Pentaceraster decipiens (Bell, 1884)
Pentaceraster gracilis (Lutken, 1871)
Pentaceraster magnificus (Goto, 1914)
Pentaceraster multispinus (van Martens, 1866)
2
3

Depth range m.
6

..

+
4+
+

4+
+
4-

..
..

+
+

4-

..

.
.

+
+
+

+
+

+
+
+
+

+
+

+
+

+
+
+
+
+

+
+
+

+
+
+
+
+
+

174-204
275
120-250
180-920
50-348
40-906
143-178
0-260
18
0-60
intertidal
0-46
0-20
30-50

+
+

156-300
686-1010
170-222
5-230
188-538
144-823
610-1020
632
366
shallow-450
134-450
115-398
172-363
219
186-+65
170-396
592-1980
170-1800

+
+
+

0-40
0-90
25-123
110-146
0-37
5-81
10-54
0-36
0-<20
3-90
0-68

+
+

+
+
+
+
+
+

0-54
40-60
0-20
0-73
10
0-27

possible synonyms of Anthenea chinensis Gray, 1840 according to Liao & Clark, 1995 :102-103.
possible synonym of Goniodiscaster scaber (Mobius, 1859) according to A.M.C. in C1ark & Rowe, 1971:
50.

472

THE RAFFLES BULLETIN OF ZOOLOGY 2000 Supplement No. 8

Pentaceraster regulus (Muller & Troschel, 1842)


Pentaceraster sibogae Diiderlein, 1916
Pentaster obtusatus (Bory de SI. Vincent, 1827)
Poraster superbus (Moebius, 1859)
Protoreaster nodosus (Linnaeus, 1758)

0-60
32-59
0-25
18-63
0-38

+
+

0-123
18-81

..

20-73

+
+
+
+

+
+

Chaetasteridae
Chaetaster moorei Bell, 1894

+
+
+

Asterodiscididae
Asterodiscides elegans (Gray, 1847)
Asterodiscides helonotus (Fisher, 1913)

Ophidiasteridae
Andora wilsoni Rowe, 1977
Bunaster ritteri Diiderlein. 1896
Celerina heffernani (Livingstone, 1931)
Dactylosaster cylindricus (Lamarck, 1816)
Fromia eusticha Fisher, 1913
Fromia hadracantha H.L. Clark, 1921
Fromia indica (Perrier, 1869)
Fromia milleporella (Lamarck, 1816)
Fromia monilis Perrier, 1875
Fromia pacifica H.L. Clark, 1921
Gomophia egyptiaca egeriae A.M. Clark, 1967
Gomophia gomophia (Perrier, 1875)
Gomophia mamillifera (Livingstone, 1930)
Heteronardoa carinata (Koehler, 1910)
Heteronardoa diamantinae Rowe, 1976
Leiaster coriaceus (Peters, 1852)
Leiaster leachi (Gray, 1840)
Leiaster speciosus von Martens, 1866
Linckia guildingi Gray, 1840
Linckia laevigata (Linnaeus, 1758)
Linckia multifora (Lamarck, 1816)
Nardoa frianti Koehler, 1910
Nardoa galatheae (Lutken, 1865)
Nardoa novaecaledoniae (Perrier, 1875)
Nardoa tuberculata Gray, 1840
Nardoa tumulosa Fisher, 19174
Neoferdina cumingi (Gray, 1840)
Neoferdina offreti (Koehler, 1910)
Ophidiaster armatus Koehler, 1910
Ophidiaster chinensis Perrier, 18755
Ophidiaster cribrarius Lutken, 1871
Ophidiaster duncani de Loriol, 1885
Ophidiaster granifer Lutken, 1871
Ophidiaster hemprichi Muller & Troschel, 1842
Ophidiaster multispinus Liao & A.M. Clark, 1996
Tamaria dubiosa (Koehler, 1910)
Tamaria jusca Gray, 1840
Tamaria megaloplax (Bell, 1884)
Tamaria pusilla (Mii11er & Troschel, 1844)

Depth range m.

Distribution

Taxa

+
+
+
+

+
L

+
+
+
+
+

+
+
+
+

+
+
+
+
+

..

+
+

+
+

+
+
+
+
+
+
+
+
+

+
+
+
+
+
+
+

...

+
+

+
+

+
+
+

+
+
+
+
+

+
+

+
+
+
+
+

+
+

+
+

..

+
+

..

18
40-54
0-55
0-11
0-55
0-20
0-44
0-73
0-51
0-30
0-80
0-40
0-16
34-230
15-187
0-52
0-183
0-81
0-46
0-60
0-69
0-51
0-25
0-5
0-70
0-62
0-30
0-62
5-450
0-31
0-20
0-123
0-45
15-55
127-757
0-70
0-117

possibly misidentified ; some SCS records believed to represent a new, unpublished species of Nardoa.
validity uncertain -see Liao & Clark, 1995: 121.

473

Lane et al.: Echinoderm fauna of the South China Sea


Distribution

Taxa

Mithrodiidae
Mithrodia clavigera (Lamarck, 1816)
Thromidia catalai, Pope & Rowe, 19776
Asteropseidae
Asteropsis carinijera (Lamarck, 1816)
Valvaster striatus (Lamarck, 1816)
Acanthasteridae
Acanthaster planci (Linnaeus, 1758)

0-81
10-105
0-55
0-25

+
+

Poraniidae
Marginaster paucispinus Fisher, 1913
Asterinidae
Anseropoda diaphana (Sladen, 1889)
Anseropoda fisheri Aziz & Jangoux, 1985
Anseropoda rosacea (Larnarck, 1816)
Asterina anomala H.L. C1ark, 1921
Asterina cepheus (Miil1er & Troschel, 1842)
Asterina coronata van Martens, 1866
Asterina limboonkengi Smith, 1927
Asterina orthodon Fisher, 1922 7
Asterina sarasini (de Lorio1, 1897)
Disasterina odontacantha Liao, 1980
Nepanthia belcheri (Perrier, 1875)
Nepanthia briareus (Bel1, 1894)
Nepanthia fisheri Rowe & Marsh, 1982
Nepanthia maculata Gray, 1840
Paranepanthia pedicellaris (Fisher, 1913)
Patiriella pseudoexigua Dartnal1, 1971
Pseudasterina delicata Aziz & Jangoux, 1985
Pseudasterina granulosa Aziz & Jangoux, 1985
Tegulaster ceylanica (Doderlein, 1889)

Deptb range m.

+
+
+
+

+
+
+

+
+
+
+

+
+

+
+

183

275
99-137
0-300

1-20
0-18

+
+

+
+
+
+

+
+
+
+

Leilasteridae
Lei/aster spinulosus Aziz & Jangoux, 1985

0-54

+
+

+
+

0-2
0-1
0-128
27-83
29-198
0-731
123
0-45
192-275
130-137
20
130-220

Order VELATIDA
Solasteridae
Seriaster regularis Jangoux, 1984
Crossaster sp.

20-65

+
+

Pterasteridae
Euretaster insignis (Sladen, 1882)

+
+
+

0-132

+
+
+
+

+
+
+
+

+
+
+

0-62
0-73
14-757
0-150

Order SPINULOSIDA
Echinasteridae
Echinaster callosus van Marenzel1er, 1895
Echinaster luzonicus (Gray, 1840)
Echinaster stereosomus Fisher, 1913
Metrodira subulata Gray, 1840
6
7

verbal report (to D.J.W.L.) of this unmistakable species at Layang Layang, 725'N;I1350'E.
possible synonym of Asterina sarasini (de Loriol, 1897) - see Clark & Rowe, 1971: 68; Liao & Clark, 1995:
131.

474

THE RAFFLES BULLETIN OF ZOOLOGY 2000 Supplement No. 8


Taxa

Distribution
2

+
+
+

+
+
+

Depth range m.
6

Order FORCIPULATIDA
Zoroasteridae
Pholidaster squamatus Sladen, 1889
Zoroaster carinatus Alcock, 1893
Zoroaster ophiactis Fisher, 1916
Asteriidae
Asterias versicolor Sladen, 1889
Coronaster halicepus Fisher, 1917
Coronaster sakuranus Doderlein, 1902
Coronaster volsellatus (Sladen, 1889)
Sclerasterias mazophorus
Wood-Mason & Alcock, 1891
Stolasterias acutispina (Stimpson, 1862)
Tarsaster stoichodes Sladen, 1889

+
+
+

..

100-399
170-1105
959-1628
0-92
190-545
192-595
59-630
150-4596

+
+

85-1224

Order BRISINGIDA
Brisingidae
Astrostephane acanthogenys (Fisher, 1916)
Brisinga sp.
Craterobrisinga analoga Fisher, 1919
Craterobrisinga eucoryne Fisher, 1916
Novodinia penichra (Fisher, 1916)
Novodinia radiata Aziz & Jangoux, 1985
Stegnobrisinga placoderma (Fisher, 1916)

to

315

....

40-659
686-1358
168-188
215-216
960-1022

to

1920

to

Freyellidae
Freyella echinata Sladen, 1889
CRINOIDEA
OroerBOURGUETICRINIDA
Bathycrinidae
Democrinus japonicus Gis1en, 1927

156-210

Order ISOCRINIDA
Isocrinidae
Diplocrinus alternicirrus Carpenter, 1884
Hypalocrinus naresianus (Carpenter, 1884)
Metacrinus interruptus Carpenter, 1884
Metacrinus multisegmentatus
Chang & Liao, 1963
Metacrinus musorstomae Roux, 1981
Metacrinus nodosus Carpenter, 1884
Metacrinus rotundus Carpenter, 1884
Metacrinus serratus Doderlein, 1907
Saracrinus angulatus (Carpenter, 1884)
Saracrinus superbus (Carpenter, 1884)

+
+
+
+

+
to

104-230
170-216
570-1152
100-500
326-550
192-230
188-310

+
+
+
+

35-56
0-59
0-c.300
0-135
<20
8-92

to

+
+

+
+
+
+

+
+
+
+
+

914-1097
686-1264
126-210

Order COMATULIDA
Comasteridae
Capillaster asterias A.H. Clark, 1931
Capillaster macrobrachius (Hartlaub, 1890)
Capillaster multiradiatus (Linnaeus, 1758)
Capillaster sentosus (Carpenter, 1888)
Comanthina audax Rowe et aI., 1986
Comanthina nobilis (Carpenter, 1888)

+
+

+
+
+
+

475

+
+

Lane et al.: Echinoderm fauna of the South China Sea


Taxa

Comanthina schlegelii (Carpenter, 1881)


Comanthus altemans (Carpenter, 1881)
Comanthus briareus (Bell. 1882)
Comanthus delicata (A. H. Clark, 1909)
Comanthus gisleni Rowe et aI., 1986
Comanthus parvicirrus (Miiller, 1841)
Comanthus wahlbergii (Miiller, 1843)
Comaster brevicirrus (Bell, 1894)
Comaster distinctus (Carpenter, 1888)
Comaster gracilis (Hartlaub, 1890)
Comaster minima (A.H. Clark, 1909)
Comaster multibrachiatus (Carpenter, 1888)
Comaster multifidus (Miiller, 1841)
Comaster teneUus A.H. Clark, 1931
Comatella maculata (Carpenter, 1888)
ComateUa nigra (Carpenter, 1888)
Comatella stelligera (Carpenter, 1888)
Comatula micraster A.H. Clark, 1909
Comatula pectinata (Linnaeus, 1758)
Comatula purpurea (Miiller, 1843)8
Comatula solaris Larnarck, 1816
Comissia littoralis (A.H. Clark, 1912)
Comissia peregrina (Bell, 1894)
Oxycomanthus bennetti (Miiller, 1841)
Oxycomanthus comanthipinna (Gislen, 1922)
Oxycomanthus grandicalyx (Carpenter, 1882)
Oxycomanthus intermedia (A.H. Clark, 1916)
Oxycomanthus japonicus (Miiller, 1841)

Zygometridae
Catoptometra hartlaubi (A.H. Clark, 1907)
Catoptometra magnifica A.H. Clark, 1908
Catoptometra rubroflava (A.H. Clark, 1907)
Zygometra comata A.H. Clark, 1911

+
+
+
+

+
+
+

+
+
+

+
+
+

+
+
+
+

+
+

+
+
+
+

+
+

+
+

+
+
+
+
+
+
+

+
+

+
+
+

+
+

+
+
+
+
+

+
+

+
+
+

+
+
+

+
+

+
+

+
+
+

+
+

+
+

+
+
+
+
+

+
+

+
+

+
+
+

+
+

+
+
+
+
+
+
+

Eudiocrinidae
Eudiocrinus indivisus (Semper, 1868)
Eudiocrinus tenuissimus Gislen, 1940
Eudiocrinus venustulus A.H. Clark, 1912
Himerometridae
Amphimetra laevipinna (Carpenter, 1882)
Amphimetra tessellata (Miiller, 1841)
Craspedometra acuticirra (Carpenter, 1882)
Heterometra africana (A.H. Clark, 1911)
Heterometra bengalensis (Hartlaub, 1890)
Heterometra crenulata (Carpenter, 1882)
Heterometra propinqua (A.H. Clark, 1912)
Heterometra pulchra A.H. Clark, 1912
Heterometra savignii (Miiller, 1841)
Heterometra variipinna (Carpenter, 1882)
Himerometra magnipinna A.H. Clark, 1908
Himerometra martensi (Hartlaub, 1890)
Himerometra robustipinna (Carpenter, 1881)

Depth range m.

Distribution

+
+
+
+

0-150
0-256

+
+

+
+
+

+
+
+

+
+
+

0-278
0-90
1-120
10-257
0-32
1-110
1-103
0-52
15-290
4-50
9-216
20-83
0-91
0-90
0-15
0-160
0-210
22-109
0-73
0-120
0-109
<20
100-110
0-50
1-20

137-278
36-914
25-183
0-150
36-183
<20
73-174
0-60
0-109
0-88
0-44
0-111
82-88
27-73(?89)
0-18
0-20
0-38
2
0-57

considered a synonym of Comatula pectinata (Linnaeus, 1758) by F.W.E.R. in Rowe & Gates, 1995: 148,
but readily distinguishable from pectinata on West Australia coast according to L.M.M.

476

THE RAFFLES BULLETIN OF ZOOLOGY 2000 Supplement No. 8


Taxa

Distribution
2

Mariametridae
Dichrometra bimaculata (Carpenter, 1881)
Dichrometra doederleini (de Lariol, 1900)
Dichrometra flagellata (Muller, 1841)
Lamprometra palmata (Muller, 1841)
Liparometra articulata (Muller, 1849)
Liparometra regalis (Carpenter, 1888)
Mariametra subcarinata (A.H. Clark, 1908)
Mariametra vicaria (Bell, 1894)
Oxymetra erinaceus (Hartlaub, 1890)
Oxymetra finschi (Hartlaub, 1890)
Stephanometra echinus (A.H. Clark, 1908)9
Stephanometra indica (Smith, 1876)
Stephanometra oxyacantha (Hartlaub, 1850)
Stephanometra spinipinna (Hartlaub, 1890)
Stephanometra tenuipinna (Hartlaub, 1890)
Colobometridae
Basilometra boschmai A.H. Clark, 1936
Cenometra bella (Hartlaub, 1890)
Colobometra discolor A.H. Clark, 1909
Colobometra perspinosa (Carpenter, 1881)
Cyllometra manca (Carpenter, 1888)
Decametra laevipinna (A.H. Clark, 1912)
Decametra mylitta A.H. Clark, 1912
Decametra parva (A.H. Clark, 1912)
Iconometra bel/ona (A.H. Clark, 1920)
Iconometra speciosa A.H. Clark, 1929
Oligometra chinensis (A.H. Clark, 1918)
Oligometra serripinna (Carpenter, 1881)
Pontiometra andersoni (Carpenter, 1888)
Tropiometridae
Tropiometra afra afra (Hartlaub, 1890)
Tropiometra afra macrodiscus (Hara, 1895)
Calometridae
Gephyrometra versicolor (A.H. Clark, 1907)
Neometra aleeto (A.H. Clark, 1911)
Neometra multicolor (A.H. Clark, 1907)
Asterometridae
Asterometra anthus (A.H. Clark, 1907)
Asterometra cristata A.H. Clark, 1911
Asterometra mirifica A.H. Clark, 1909
Pterometra pulcherrima (A.H. Clark, 1909)
Pterometra trichopoda (A.H. Clark, 1908)
Pterometra venusta A.H. Clark, 1912

+
+
+

7
0-1
0-150
0-45
0-50
0-69
3-31
40-108
53-100
0-50
0-77
0-38
0-73
0-27
0-245
0-48

+
+
+

+
+
+
+

+
+

...

+
+

+
+

+
+
+

+
+

+
+

+
+
+
+
+
+

+
+

+
+

0-103
0-55
42-106
0-122
22-329
0-3
25-85
1-90
16-73
146
0-2
0-90
0-73

+
+

0-110
0-66

+
+

5-174
77-208
20-600

+
+

...

+
+

+
+

+
+
+
+

...

+
+
+
+

...

+
+

+
+
+
+
+
+

+
+

Thalassometridae
Parametra orion (A.H. Clark, 1907)
Antedonidae
Antedon parviflora (A.H. Clark, 1912)
Antedon serrata A.H. Clark, 1908
Dorometra aphrodite (A.H. Clark, 1912)10
Dorometra parvicirra (Carpenter, 1888)

Depth range m.

+
+
+
+

+
+
+
+

64-210
80-146
73-183
36-174
68-140
5-80
128-306
0-275
0-180
48-174
0-164

a probable synonym of Stephanometra tenuipinna (Hartlaub, 1890) (see Clatk & Rowe, 1971: 24), but genus
Stephanometra is in need of revision.
10 close to, possibly synonymous with, Dorometra parvicirra (Carpenter, 1888) - see Liao & Clatk, 1995 : 62.

477

Lane et al.: Echinoderm fauna of the South China Sea


Taxa

Distribution
2

Dorometra nana (Hartlaub, 1890)


Euantedon sinensis A.H. Clark, 1912
Mastigometra pacifica A.H. C1ark, 1918
Toxometra paupera A.H. C1ark, 1911

Depth rauge m.
6

7
0-60
deep
littoral
0-510

+
+
+
+

OPHIUROIDEA
Order PHRYNOPHIURlDA
Ophiomyxidae
Astrogymnotes catastica H.L. Clark, 1914
Neoplax crassipes Koehler, 1922
Ophiobyrsella intorta Koehler, 1922
Ophiodera neglecta Koehler, 1904
Ophiomyxa australis Liitken, 1869
Ophiomyxa bengalensis Koehler, 1897
Ophiomyxa longipeda Brock, 1888
Ophiophrixus confinis Koehler, 1922
Euryalidae
Astroceras mammosus Koehler, 1930
Astroceras pergamena Lyman, 1879
Euryale aspera Lamarck, 1816
Euryale purpurea Mortensen, 1934
Sthenocephalus indicus Koehler, 1898
Trichaster acanthifer Doderlein, 1927
Trichaster palmiferus (Lamarck, 1816)
Asteronychidae
Asteronyx loveni Miiller & Troschel, 1842
Astrodia tenuispina (Verrill, 1885)
Gorgonocephalidae
Asteroporpa hadracantha H.L. Clark, 1911
Astroboa nuda (Lyman, 1874)
Astrocladus dofleini Doderlein, 1911
Astrocladus exiguus (Lamarck, 1816)
Astrocladus tonganus DOderlein, 1911
Astroclon propugnatoris Lyman, 1879
Astroglymna sculptum (DOderlein, 1896)
Astrotoma drachi Guille, 1981
Astrotoma manilense Doderlein, 1927
Gorgonocephalus dolichodactylus Doderlein, 1911

40-205
24
1346
65-744
0-1006
296-1962

448-558

>I-

+
+
+

+
+

+
+
+
+
+
+

+
+
+
+

+
+

+
+

+
+

+
+
+

+
+
+
+

+
+
+

+
+
+

...

+
+

90-195
65-1033
0-290

36-487
18-39
33-159

109-2963
510-3720
62-194
1-113
27-278
18-494
2-10
170-236
54-300
975-1125
720
146-1134

Asteroschematidae
Asteroschema ferox Koehler, 1904

170-204

Hemieuryalidae
Ophiogyptis nodosa Koehler, 1905

35-308

Order OPHIURlDA
Ophiacanthidae
Amphilimna granulosa Liao, 1989
Amphilimna multispina Koehler, 1922
Amphilimna polyacantha Liao, 1983
Amphilimna sinica Liao, 1989
Amphilimna tanyodes Devaney, 1974
Ophiacantha composita Koehler, 1896
Ophiacantha duplex Koehler, 1897
Ophiacantha gracilis (Studer, 1882)

...

+
+
+
+

....
+
+
+

478

280
91-357
72-200
280
189
685-757
686-1234
0-400

THE RAFFLES BULLETIN OF ZOOLOGY 2000 Supplement No. 8


Distribution

Taxa
2
Ophiacantha indica Ljungman, 1867
Ophiacantha longidens Lyman, 1874
Ophiacantha pentagona Koehler, 1897
Ophialcoea congesta (Koehler, 1904)
Ophiocamax rugosa Koehler, 1904
Ophiocopa singularis Koehler, 1922
Ophiolimna perfida (Koehler, 1904)
Ophiomitra plicata Lyman, 1874
Ophiomitrella subjecta Koehler, 1922
Ophiophthalmus honestus Koehler, 1930
Ophioplinthaca globata Koehler, 1922
Ophioplinthaca hastata Koehler, 1922
Ophioplinthaca manillae Guille, 1981
Ophioplinthaca pulchra Koehler, 1904
Ophioplinthaca rudis (Koehler, 1897)
Ophiotrema tertium Koehler, 1922
Ophiotreta gratiosa (Koehler, 1897)
Ophiotreta matura (Koehler, 1904)
Ophiotreta speciosa Guille, 1981
Ophiotreta valenciennesi (Lyman, 1874)
Ophiurothamnus musorstomae Guille, 1981

Amphiuridae
Amphilycus scripta (Koehler, 1904)
Amphiodia minuta H.L. Clark, 1939
Amphiodia (Amphispina) crassa (Koehler, 1904)
Amphiodia (Amphispina) microplax Burfield, 1924
Amphiodia (Ophiophragmus) olivacea
(Brock, 1888)
Amphioplus (Amphichilus) impressus
(Ljungman, 1867)
Amphioplus (Amphichilus) intermedius
(Koehler, 1905)
Amphioplus (Amphioplus) cyrtacanthus
H.L. Clark, 1915
Amphioplus (Amphioplus) exsecratus
(Koehler, 1905)
Amphioplus (Amphioplus) iuxtus Murakami, 1943
Amphioplus (Amphioplus) legatus Koehler, 1922
Amphioplus (Amphioplus) lucidus Koehler, 1922
Amphioplus (Amphioplus) rhadinobrachius
H.L. Clark, 1911
Amphioplus (Lymanella) andreae (Ltitken, 1872)
Amphioplus (Lymanella) depressus
(Ljungman, 1867)
Amphioplus (Lymanella) laevis (Lyman, 1874)
Amphipholis loripes Koehler, 1922
Amphipholis misera (Koehler, 1899)
Amphipholis sobrina Matsumoto, 1917
Amphipholis squamata (Delle Chiaje, 1829)
Amphiura diomedeae Ltitken & Mortensen, 1899
Amphiura grandisquama Lyman, 1909
Amphiura (Amphiura) abbreviata Koehler, 1905
Amphiura (Amphiura) aestuarii Matsumoto, 1915
Amphiura (Amphiura) ambigua Koehler, 1905
Amphiura (Amphiura) bidentata H.L. Clark, 1938
Amphiura (Amphiura) crossota Murakami, 1943

Depth range m.
6

t
t
t

t
t

t
t
t

,.
t

+
+
t

t
t

+
t

+
t

+
+

<20
36-37
453-2291
5-69

t
t
t

<20-290
90-183
78-1724
450-1000
82-520
340-686
592-1125
1920
353-485
183-385
686-969
353-368
170-200
204-171
650-3124
1335
50-925
239-4161
194-200
245-385
975-1125

t
t

8-220

10-105

<20
<20
721
3-152

+
t

16-194

291

t
t
t

t
t
t

t
t

t
t
t
t
t
t
t

479

6-160
7-100
78-196
18-348
137-550
0-500
368-1125
10-686
13-40
125-230
0-34
0-37

Lane et al.: Echinoderm fauna of the South China Sea


Distribution

Taxa
2

Amphiura (Amphiura) divaricata Ljungman,1867


Amphiura (Amphiura) duncani Lyman, 1882
Amphiura (Fellaria) ecnomiotata H.L. Clark, 1911
Amphiura (Fellaria) vadicola Matsumoto, 1915
Amphiura (Ophiopeltis) phalerata (Lyman, 1874)
Amphiura (Ophiopeltis) tenuis (H.L. Clark, 1938)
Dougaloplus acanthinus (H.L. Clark, 1911)
+
Dougaloplus echinatus (Ljungman, 1867)
+
Ophiocentrus anomalus Liao, 1983
Ophiocentrus aspera (Koehler, 1905)
Ophiocentrus dilatatus (Koehler, 1905)
Ophiocentrus inaequalis (H.L. Clark, 1915)
Ophiocentrus koehleri Gislen, 1926
Ophiocentrus putnami (Lyman, 1871)
Ophionephthys difficilis (Duncan, 1887)
Ophiostigma rugosum H.L. Clark, 1918
Paracrocnida sinensis (A.H. Clark, 1917)
Paramphichondrius tetradontus
Guille & Wolff, 1984
Ophiactidae
Histampica duplicata (Lyman, 1875)
Ophiactis affinis Duncan, 1879
Ophiactis definita Koehler, 1922
Ophiactis fuscolineata H.L. Clark, 1938
Ophiactis hexacantha H.L. Clark, 1939
Ophiactis macrolepidota
Marktanner-Turneretscher, 1887
Ophiactis maculosa von Martens, 1870 11
Ophiactis modesta Brock, 1888 11
Ophiactis picteti (de Loriol, 1893)
Ophiactis profundi Liitken & Mortensen, 1899
Ophiactis quadrispina H.L. Clark, 1915 11
Ophiactis savignyi (Miiller & Troschel, J842)
Ophiactis versicolor H.L. Clark, 1939 11
Ophiodaphne formatus (Koehler, 1905)
Ophiodaphne materna Koehler, 1930
Ophiosphaera insignis Brock, 1888

+
+

+
+
+
+

+
+
+
+

+
+

+
+
+

Depth range m.

+
+
+
+

....

+
+

..

35-164

+
+

+
+

+
+
+

+
+
+

+
+

+
+

+
+

+
+

+
+

Ophiotrichidae
Gymnolophus obscura (Ljungman, 1867)
Macrophiothrix coerulea (Djakonov, 1930)
Macrophiothrix demessa (Lyman, 1861)
Macrophiothrix expedita (Koehler, 1905)
Macrophiothrix fumaria
(Miiller & Troschel, 1842)12
Macrophiothrix galateae (Liitken, 1872)
+
Macrophiothrix hirsuta (Miiller & Troschel, 1842)
Macrophiothrix hybrida (H.L. Clark, 1915)12
Macrophiothrix koehleri A.M. Clark, 1968
Macrophiothrix lineocaerulea (H.L. Clark, 1928)12
Macrophiothrix longipeda (Lamarck, 1816)
Macrophiothrix lorioli A.M. Clark, 1968
Macrophiothrix martensi (Lyman, 1874)13

+
+
+
+
+
+
+

+
+

.
+
+

+
+

+
+
+

*
+
+

+
+
+
+
+
+
+
+

+
+

+
+
+
+
+
+
+
+

7-78
3-400
58-290
0-c.30
<20
0-12
10-85
16-118
19-62
0-36
216
18-36
30-84
<20-90
I-50

+
+
+

685-1125
0-90
250-1280
0-15
0-183
1-177
<20
0-180
1-50
46-410
0-1000
<20-29
5-274
9-249
18-245
0-52
1-20
0-128
0-73
<20
30-50
0-90
47
0-102
littoral-lOO
0-290
0-24
0-73

11 these 4 spp. considered to be synonymous with Ophiactis savignyi (MUller & Troschel, 1842) by Cherbonnier
& Guille (1978) but not according to Liao & Clark, 1995:218.
12 transferred from subgenus Ophiothrix (Placophiothrix) - see Hoggett, 1992; Price & Rowe, 1996: 75-76.
13 transferred from subgenus Ophiothrix (Keystonea) - see Hoggett, 1992; Price & Rowe, 1996: 75-76.

480

THE RAFFLES BULLETIN OF ZOOLOGY 2000 Supplement No. 8

Macrophiothrix melanosticta (Grube, 1868)"


Macrophiothrix nereidina (Lamarck, 1816)13
Macrophiothrix propinqua (Lyman, 1861)
Macrophiothrix robillardi (de Loriol, 1893)
Macrophiothrix striolata (Grube, 1868)12
Macrophiothrix variabilis (Duncan, 1887)
Macrophiothrix vicina (Koeh1er, 1930)13
Macrophiothrix virgata (Lyman, 1861)12
Ophiocnemis marmorata (Lamarck, 1816)
Ophiogymna elegans Ljungman, 1867
Ophiogymna funesta Koehler, 1922
Ophiogymna pellicula (Duncan, 1876)
Ophiogymna pulchella (Koehler, 1905)
Ophiomaza cacaotica Lyman, 1871
Ophiopsammium semperi Lyman, 1874
Ophiopteron elegans Ludwig, 1888
Ophiopteron vitiense Koehler, 1927
Ophiothela danae Verrill, 1869
Ophiothela venusta (de Loriol, 1900)
Ophiothrix capillaris Lyman, 1879
Ophiothrix contenta Koehler, 1930
Ophiothrix eurycolpodes H.L. Clark, 1918
Ophiothrix (Acanthophiothrix) deceptor
Koehler, 1922
Ophiothrix (Acanthophiothrix) leucotrigona
H.L. Clark, 1918
Ophiothrix (Acanthophiothrix) proteus
Koehler, 1905
Ophiothrix (Acanthophiothrix) purpurea
von Martens, 1867
Ophiothrix (Acanthophiothrix) scotiosa
Murakami, 1943
Ophiothrix (Acanthophiothrix) signata
Koehler, 1922
Ophiothrix (Acanthophiothrix) spinosissima
Koehler, 1905
Ophiothrix (Acanthophiothrix) vetusta
Koehler, 1930
Ophiothrix (Acanthophiothrix) vigelandi
A.M. Clark, 1968
Ophiothrix (Acanthophiothrix) viridialba
von Martens, 1867
Ophiothrix (Ophiothrix) ciliaris (Lamarck, 1816)
Ophiothrix (Ophiothrix) crassispina Koehler, 1904
Ophiothrix (Ophiothrix) elegans Liitken, 1869
Ophiothrix (Ophiothrix) exigua Lyman, 1874
Ophiothrix (Ophiothrix) foveolata
Marktanner-Tumeretscher, 1887
Ophiothrix (Ophiothrix) infirma Koehler, 1905
Ophiothrix (Ophiothrix) koreana Duncan, 1879
Ophiothrix (Ophiothrix) marenzelleri Koehler, 1904
Ophiothrix (Ophiothrix) miles Koehler, 1905
Ophiothrix (Ophiothrix) plana Lyman, 1874
Ophiothrix (Ophiothrix) prostata Koehler, 1922
Ophiothrix (Ophiothrix) rotata von Martens, 1870

Depth range m.

Distribution

Taxa

+
+

+
+
+
+
+
+

+
+

+
+
+

+
+
+
+
+
+
+

+
+

+
+
+

+
+

et

+
+
+

+
+
+

+
+
+
+

+
+
+
+

+
+

+
+
+

+
+

..

+
+

0-82
0-73
0-80
-5
0-89
0-139
10
0-100
1-348
80-300
10-116
73-247
0-80
0-50
0-189
25-50
0-120
0-40
22-750

<to

+
+

96-107

<to

0-125

+
+

+
+

+
129-139

+
+

2-36

191-245

+
+

0-618

+
+

+
+

+
+

..

+
+

+
+

+
+

+
+

+
+

<to

481

0-1046

11-170
0-308
186-274
0-67
0-140
0-141
54-200
3-400
0-128
0-14
0-70
2-46
9

Lane et al.: Echinoderm fauna of the South China Sea


Distribution

Taxa

Ophiothrix (Ophiothrix) savignyi


(MUller & Troschel, 1842)
Ophiothrix (Ophiothrix) trilineata LUtken, 1869
Ophiothrix (Ophiothrix) vitrea DOderlein, 1896
Ophiothrix (Theophrix) pusilla Lyman, 1874
Ophiocomidae
Ophiarthrum elegans Peters, 1851
Ophiarthrum pietum (MUller & Troschel, 1842)
Ophioeoma anaglyptiea Ely, 1944
Ophioeoma brevipes Peters, 1851
Ophioeoma dentata MUller & Troschel, 1842
Ophioeoma erinaeeus MUller & Troschel, 1842
Ophioeoma pica MUller & Troschel, 1842
Ophioeoma pusilla (Brock, 1888)
Ophioeoma sehoenleini MUller & Troschel, 1842
Ophioeoma seolopendrina (Lamarck, 1816)
Ophioeomella sexradia (Duncan, 1887)
+
Ophiomastix annulosa (Lamarck, 1816)
Ophiomastix asperula LUtken, 1869
Ophiomastix earyophyllata LUtken, 1869
Ophiomastix eorallieola H. L. Clark, 1915
Ophiomastix flaeeida Lyman, 1874
Ophiomastix janualis Lyman, 1871
Ophiomastix mixta LUtken, 1869
Ophiomastix variabilis Koehler, 1905
Ophiomastix venosa Peters, 1851
Ophiopsila abscissa Liao, 1982
Ophiopsila pantherina Koehler, 1898
Ophionereididae
Ophioehiton jastigatus Lyman, 1878
Ophionereis dubia amoyensis A.M. Clark, 1953
Ophionereis dubia dubia (MUller & Troschel, 1842) .
Ophionereis jusea Brock, 1888
Ophionereis porreeta Lyman, 1860
Ophionereis variegata Duncan, 1879
Ophiodermatidae
Bathypeetinura heros (Lyman, 1879)
Cryptopelta longibraehialis Koehler, 1930 14
Ophiarachna affinis LUtken, 1869
Ophiaraehna inerassata (Lamarck, 1816)
Ophiaraehna ohshimai Murakami, 1943
Ophiarachnella elegans (Bell, 1894)
Ophiaraehnella gorgonia (MUller & Troschel, 1842) .
Ophiaraehnella injemalis
(MUller & Troschel, 1842)
+
Ophiaraehnella maeraeantha H.L. Clark, 1909
Ophiaraehnella paueispina (Koehler, 1905)
Ophiaraehnella septemspinosa
(MUller & Troschel, 1842)
Ophiaraehnella similis (Koehler, 1905)
Ophiaraehnella stabilis (Koehler, 1905)
Ophioehaeta hirsuta LUtken, 1869
Ophioch'asma stellatum (Ljungman, 1867)
+
Ophioeonis cineta Brock, 1888

+
+
+

+
+

*
+
+
+
+
+
+

+
+
+
+
+

+
+
+

+
+

...

+
+
+

+
+
+
+
+
+

+
+
+
+
+
+

+
+
+
+
+

+
+
+
+
+
+
+
+

+
+
+
+

+
+

+
+

+
+
+

+
+
+

+
+
+
+
+
+
+
+

*
+
+
+

Depth range m.

+
+

24-64
0-50

+
+

+
+

+
+
+

+
+

128-1628
0-34
0-230
0-45
0-60
0-60

+
+

...
+

30-120
0-100

+
+

+
+

0-36
0-20
0-20
0-54
0-35
0-27
0-24
0-20
0-20
0-13
0-15
0-57
0-20
0-54
0-20
0-82
0-22
0-54
0-60

240-2960
13-243
0-25
0-97

...

0-54
3-300
5-54

...

+
+

0-233
30
34-83
0-55
0-59
12-83
0-26
0-110
0-111

14 possible synonym of Cryptopelta granulifera H.L. Clark, 1909; see Rowe & Gates, 1995: 395.

482

THE RAFFLES BULLETIN OF ZOOLOGY 2000 Supplement No. 8


Taxa
2

Ophioconis cupida Koehler, 1905


Ophioderma tonganum Liitken, 1872 15
Ophiodyscrita acosmeta H. L. Clark, 1938
Ophiopeza fallax Peters, 1851
Ophiopeza spinosa (Ljungman, 1867)
Ophiopsammus aequalis (Lyman, 1880)
Ophiopsammus yoldii (Liitken, 1856)
Ophioleucidae
Ophiocirce inutilus Koehler, 1904
Ophioleuce seminudum Koehler, 1904
Ophiopallas paradoxa Koehler, 1904
Ophiuridae
Amphiophiura canaliculata Koehler, 1922
Amphiophiura insolita (Koehler, 1904)
Amphiophiura paupera (Koehler, 1896)
Amphiophiura pertusa Koehler, 1930
Amphiophiura spatulijera Koehler, 1922
Amphiophiura sculptilis (Lyman, 1878)
Amphiophiura sordida (Koehler, 1897)
Ophiolepis cardioplax Murakami, 1943
Ophiolepis cineta Miiller & Troschel, 1842
Ophiolepis rugosa Koeh1er, 1898
Ophiolepis superba H.L. Clark, 1915
Ophiolipus granulatus Koehler, 1897
Ophiomastus tegulitius Lyman, 1878
Ophiomusium corticosum Lyman, 1878
Ophiomusium facetum Koehler, 1922
Ophiomusium facundum Koehler, 1922
Ophiomusium lymani Thomson, 1873
Ophiomusium luetkeni Lyman, 1878
Ophiomusium sealare Lyman, 1878
Ophiomusium simplex Lyman, 1878
Ophioplocus declinans (Koehler, 1904)
Ophioplocus imbricatus (Miiller & Troschel, 1842)
Ophioplocus japonicus H. L. Clark, 1911
Ophiotylos brevipes Liao, 1978
Ophiotylos leucus Murakami, 1943
Ophiozonella bispinosa (Koehler, 1897)
Ophiozonella molesta (Koehler, 1904)
Ophiozonella subtilis Koehler, 1922
Ophiura aequalis Lyman, 1878
Ophiura clemens (Koehler, 1904)
Ophiura flagellata (Lyman, 1878)
Ophiura kinbergi (Ljungman, 1867)
+
Ophiura lanceolata H.L. Clark, 1939
Ophiura micracantha H.L. Clark, 1911
Ophiura pteracantha Liao, 1983
Sinophiura multispina (Koehler, 1922)
Stegophiura hainanensis Liao, 1995
Stegophiura sterilis Koeh1er, 1922

Depth range m.

Distribution
3

+
+

10-600

0-20
25
0-74
161-401
0-215

+
+
+

+
+
+
+
+

+
+

...
+
+
+

+
+

...

+
+
+
+
+
+
+

+
+
+
+

...
...
+
+

+
+

+
+
+

+
+

+
+

+
+
+
+

+
+

+
+

+
+

+
+

...

112-209
50-1635
112-500
106
204-1236
743-1289
179-300
959
457-3429
250-1280
littoral
0-45
5-73
0-30
205-470
510-4840
3383
513-1752
454-1400
130-4000
196-1125
124-1224
100-300
183-187
0-30
0-2
0-5

+
+
+
+
+

...

+
+
+
+
+
+
+

+
+

+
+
+
+
+
+

...
...

+
+
+
+
+

150-382
68-1270
149-929
694-2194
686-1264
96-2330
0-500
122-293
144-472
3-93
51-357
158
380

15 presence of genus Ophioderma in lndo-west Pacific questionable - see comments by A.M.C. in Clark &
Rowe, 1971: 126.

483

Lane et al.: Echinoderm fauna of the South China Sea


Taxa

Distribution
2

Depth range m.
6

+
+
+
+
+

ECHINOIDEA
Order CIDAROIDA

Cidaridae
Chondrocidaris brevispina H.L. Clark, 1925
Eucidaris metularia (Lamarck, 1816)
Goniocidaris tenuispina Mortensen, 1927
Phyllacanthus imperialis (Lamarck, 1816)
Prionocidaris australis (Ramsay, 1885)
Prionocidaris baculosa (Lamarck, 1816)
Prionocidaris bispinosa (Lamarck, 1816)
Prionocidaris verticillata (Lamarck, 1816)
Stereocidaris grandis Diiderlein, 1887
Stereocidaris indica Diiderlein, 1901
Stylocidaris annulosa Mortensen, 1927
Stylocidaris reini (Diiderlein, 1887)

+
+
+

+
+

0-15
0-570
570-1411
0-73
10-145
0-250
4-125
0-54
204-716
400-1298
125-300
100-841

70-540

+
+

+
+

+
+

+
+
+
+
+

+
+
+

+
+

+
+
+

...

OmerECHINOTHUmOIDA

Echinothuriidae
Araeosoma owstoni Mortensen, 1904
Araeosoma tesselatum var. carinatum
Mortensen, 1934
Asthenosoma heteractis Bedford, 1900
Asthenosoma ijimai Yoshiwara, 1897 16
Asthenosoma varium Grube, 1868
Calveriosoma gracile (A. Agassiz, 1881)
Hygrosoma hoplacantha (Thomson, 1877)

...
+

...

+
L

Phormosomatidae
Phormosoma bursarium A. Agassiz, 1881

+
+
+
+

+
+

385
9
7-300
0-167
145-950
360-2068
170-2340

Order DIADEMATOIDA

Aspidodiadematidae
Aspidodiadema tonsum A. Agassiz, 1879
Plesiodiadema indicum (Diiderlein, 1900)
Diadematidae
Astropyga radiata (Leske, 1778)
Chaetodiadema granulatum Mortensen, 1903
Chaetodiadema japonicum Mortensen, 1904
Diadema paucispinum A. Agassiz, 1863
Diadema savignyi Michelin, 1845
Diadema setosum (Leske, 1778)
Echinothrix calamaris (pallas, 1774)
Echinothrix diadema (Linnaeus, 1758)
Eremopyga debilis Mortensen, 1940
Eremopyga denudata (de Meijere, 1904)

180-1135
240-1150

+
+

+
+

+
+
+
+

+
+
+
+
+
+

...

+
*
+
+
+
+

+
+
+
+

+
+
+
+
+

+
+

+
+

+
+
+
+

+
+

0-70
15-216
50-135
0-40
0-70
0-70
0-90
3-10
200-400
70-275

Order PHYMOSOMATOIDA

Arbaciidae
Arbacia incisa (A. Agassiz, 1863)
Coelopleurus maillardi (Michelin, 1862)
Coelopleurus maculatus
A. Agassiz & H.L. Clark, 1907
Stomopneustidae
Stomopneustes variolaris (Lamarck, 1816)

0-29
23-250

60-360

+
+

0-10

16 status of Asthenosoma ijimae Yoshiwara, 1897 uncertain - see Weinberg & de Ridder, 1998: 44.

484

THE RAFFLES BULLETIN OF ZOOLOGY 2000 Supplement No. 8


Taxa

Depth range m.

Distribution
2

Order MICROPYGOIDA

Micropygidae
Micropyga tuberculata A. Agassiz, 1879
Micropyga violacea de Meijere, 1904

+
+

150-1340
625-925

e.245-985

Order PEDINOIDA

Pedinidae
Caenopedina indica (de Meijere, 1904)
Order TEMNOPLEUROIDA

Temnopleuridae
Desmechinus anomalus H.L. Clark, 1923
Desmechinus rujus (Bell, 1894)
Desmechinus versicolor (Mortensen, 1904)
Mespilia globulus (Linnaeus, 1758)
Microcyphus ceylanicus Mortensen, 1942
Paratrema doederleini (Mortensen, 1904)
Prionechinus forbesianus (A. Agassiz 1881)
Salmaciella dussumieri (L. Agassiz, 1846)
Salmacis belli Doderlein. 1902
Salmacis bicolor L. Agassiz, 1846
Salmacis sphaeroides (Linnaeus, 1758)
Salmacis virgulata L. Agassiz, 1846
Temnopleurus alexandri (Bell, 1884)
Temnopleurus apodus
(A. Agassiz & H. L. Clark, 1907)
Temnopleurus decipiens (de Meijere, 1904)
Temnopleurus hardwicki (Gray, 1855)
Temnopleurus reevesi (Gray, 1855)
Temnopleurus toreumaticus (Leske, 1778)
Temnotrema bothryoides (L. Agassiz, 1846)
Temnotrema maculatum (Mortensen, 1904)
Temnotrema reticulatum (Mortensen, 1904)
Temnotrema sculptum A. Agassiz, 1863
Temnotrema siamense (Mortensen, 1904)
Temnotrema xishaensis Liao, 1978

..

+
+

..

+
+
+

+
+
+
+
+

+
+
+
+
+

+
+

..

+
+
+
+

+
+
+
+
+

..

+
+

+
+
+
+
+
+

+
*

+
+

+
+
+

+
+
+
+
+
+
+
+
+
+

20-80
70-100
0-e.60
<20
10-20
139-1370
7-180
10-125
0-122
0-e.90
10-55
9-73
18-500
18-40
e.5-65
e.5-565
0-82
73-80
10-112
e.25-290
0-e.500
e.5-350

Order ECHINOIDA

Toxopneustidae
Cyrtechinus verruculatus (LUtken, 1864)
Gymnechinus epistichus H.L. Clark, 1912
Goniopneustes pentagonus (A.Agassiz, 1872)
Nudechinus ambonensis Mortensen, 1942
Nudechinus inconspicuus (Mortensen, 1904)
Nudechinus multicolor (Yoshiwara, 1898)
Nudechinus stictus H.L. Clark, 1912
Pseudoboletia indiana (Miehelin, 1862)
Pseudoboletia maculata Trosehel, 1869
Toxopneustes pileolus (Lamarek, 1816)
Tripneustes gratilla (Linnaeus, 1758)
Parasaleneiidae
Parasalenia gratiosa A. Agassiz, 1863
Parasalenia poehli Pfeffer, 1887

+
+

+
+

+
+

+
+

485

+
+
+
+
+
+

0-e.130
0-63
e.50-90
0-c.50
0-20
0-10
13-20
0-100
10-82
0-90
0-e.75

+
+

0-70
0-70

+
+
+
+

..

Lane et al.: Echinodenn fauna of the South China Sea


Taxa

Distribution
2

Echinometridae
Anthocidaris crassispina (A. Agassiz, 1863)
Colobocentrotus mertensi Brandt, 1835
Colobocentrotus pedifer (de B1ainville, 1825)
Echinometra mathaei (de Blainville, 1825)
Echinometra oblonga (de B1ainville, 1825)
Echinostrephus aciculatus A. Agassiz, 1863
Echinostrephus molaris (de B1ainville, 1825)
Heterocentrotus mammillatus (Linnaeus, 1758)
Heterocentrotus trigonarius (Lamarck, 1816)

+
+

+
+
+

+
+

+
+
+

+
+
+

Strongy1ocentrotidae
Hemicentrotus pulcherrimus (A. Agassiz, 1863)

Depth range m.
6

0-70
littoral

+
+
+
+

+
+

0-139
0- ?
0-c.50
0-c.50
0-30
0-36
0-c.45

Order CASSIDULOIDA

Echino1ampadidae
Conolampas diomedeae Mortensen, 1948
Echinolampas alexandri de Lorio1, 1876
Echinolampas ovata (Leske, 1778)

+
+

+
+

181-265
c. 8-365
c. 9-75

+
+

Order HOLECTYPOIDA

Echinoneidae
Echinoneus abnormalis de Loriol, 1883
Echinoneus cyclostomus Leske, 1778
Micropetalon purpureum
A. Agassiz & H. L. Clark,1907

+
+

0-85
0-120

c.45-70

25-56

Order CLYPEASTEROIDA

C1ypeasteridae
Clypeaster (Coronanthus) latissimus
(Lamarck, 1816)
+
Clypeaster (Leptoclypus) annandalei
Koeh1er, 1922
Clypeaster (Leptoclypus) rarispinus
de Meijere, 1902
Clypeaster (Rhaphidoclypus) fervens Koeh1er, 1922
Clypeaster (Rhaphidoclypus) reticulatus
(Linnaeus, 1758)
+
Clypeaster (Stolonoclypus) humilis (Leske, 1778)
Clypeaster (Stolonoclypus) miniaceus
H.L.C1ark, 1925
Clypeaster (Stolonoclypus) virescens
DOderlein, 1885
+
Arachnoididae
Arachnoides placenta (Linnaeus, 1758)
Fibu1ariidae
Echinocyamus crispus Mazzetti, 1894
Echinocyamus provectus de Meijere, 1904
Echinocyamus scaber de Meijere, 1904
Fibularia acuta Yoshiwara, 1898
Fibularia angulipora Mortensen, 1948
Fibularia cribellum de Meijere, 1904
Fibularia oblonga Gray, 1851
Fibularia ovulum Lamarck, 1816
Fibularia volva L. Agassiz, 1847

+
+

+
+

160-475

+
+

+
+

+
+

7-91
0-216

0-125
0-216

..

c.70-230

27-630

0-57

+
+

+
+

..

+
+

+
+
+

+
+
+
+

+
+
+

+
+
+

486

+
+

+
+
+
+
+
+
+

+
+
+
+
+
+

13-564
75-390
c.200-1886
0-90
0-c.15
14-522
24
0-385
0-94

THE RAFFLES BULLETIN OF ZOOLOGY 2000 Supplement No. 8


Taxa

Laganidae
Laganum decagonale (de Blainville, 1827)
Laganum depressum Lesson, 1841
Laganum laganum (Leske, 1778)
Laganum fudsiyama Diiderlein, 1885
Peronella lesueuri (Va1enciennes, 1841)
Peronella macroproctes Koeh1er, 1922
Peronella minuta (de Meijere, 1904)
Peronella orbicularis (Leske, 1778)
Peronella pellucida Diiderlein, 1885
Peronella rubra (Doderlein, 1885)
Astriclypeidae
Astriclypeus manni Verrill, 1867
Echinodiscus auritus Leske, 1778
Echinodiscus bisperforatus Leske, 1778
Echinodiscus tenuissimus (L.Agassiz, 1847)

Distribution
2

+
+
+

+
+

+
+
+
+
+
+
+
+
+

+
+
+
+
+

+
+
+
+
+
+

+
+
+
+
+

+
+

+
+

c.5-300
0-c.85
0-34
c.50-654
0-c.70
5-106
c.l3-72
0-70
c.75-550
5-60

+
+
+

+
+
+
+

0-40
0-57
<20
0-c.20

'"

lower shore

+
+
+

+
+
+
+

+
+

+
+
+

Dendrasteridae
Sinaechinocyamus mai (Wang, 1984)
Order SPATANGOIDA

Spatangidae
Gymnopatagus magnus
A. Agassiz & H.L. Clark, 1907
Maretia cordata Mortensen, 1948
Maretia planulata (Lamarck, 1816)
Nacospatangus alta (A. Agassiz, 1863 )

+
+

+
+

Palaeostomatidae
Palaeostoma mirabile (Gray, 1851)

Asterostomatidae
Argopatagus vitreus A. Agassiz, 1879
Heterobrissus niasicus (Diiderlein, 1901)
Linopneustes excentricus de Meijere, 1902
Linopneustes fragilis (de Meijere, 1902)
Linopneustes murrayi (A. Agassiz, 1879)
Platybrissus roemeri Grube, 1866
Platybrissus ovalis (Mortensen, 1948)
Plesiozonus diomedeae Mortensen, 1948
Plesiozonus tenuis David & de Ridder, 1989

+
+

+
+
+
+

+
+
+
+

Pericosmidae
Pericosmus melanostomus Mortensen, 1948
Pericosmus porphyrocardius McNamara, 1984

+
+

+
+

Schizasteridae
Faorina chinensis Gray, 1851
Moira lachesinella Mortensen, 1930
Moira stygia A. Agassiz, 1872

+
+
+

+
+
+

c.780-1730
c.50-150
0-c.60
2-204

c.20-110

+
+

*
+
+
+
+
+
+

487

+
+
+
+

+
+
+
+

+
+

+
+

+
+
+
+
+

+
+
+
+
+

+
+

+
+

'+"

+
+
+

'"

+
+

+
+
+

+
+
+

+
+
+

Loveniidae
Echinocardium cordatum (Pennant, 1777)
Homolampas lovenioides Mortensen, 1948
Lovenia doederleini Mortensen, 1950
Lovenia elongata (Gray, 1845)
Lovenia gregalis Alcock, 1893
Lovenia subcarinata (Gray, 1845)
Lovenia triforis Koehler, 1914

Depth range m.

+
+
+

+
+

c.460-2250
125-475
694-1788
c.560-1170
620-1615
0-100
192
c.180-345
181-190
0-c.230
987
10-35
0-94
c.274-930
c.10-49
43-270
18-70
217-420
41-220
<50
6-15

Lane et al.: Echinoderm fauna of the South China Sea


Taxa
2
Proraster affinis (Mortensen, 1948)
Prymnaster rostratus (Smith, 1878)
Schizaster (Paraster) compactus (Koehler, 1914)
Schizaster (Schizaster) lacunosus (Linnaeus, 1758)
Brissidae
Anametalia sternaloides (Bolau, 1874)
Brissopsis luzonica (Gray, 1851)
Brissopsis oldhami Alcock, 1893
Brissopsis similis Mortensen, 1948
Brissus latecarinatus (Leske, 1778)
Eupatagus magnus (A. Agassiz &
H. L. Clark, 1907)
Metalia dicrana H.L. Clark, 1917
Metalia spatagus (Linnaeus, 1758)
Metalia sternalis (Lamarck, 1816)
Rhynobrissus pyramidalis A. Agassiz, 1872

Depth range m.

Distribution
3

+
+

+
+
+
+
+

+
+
+

7
915-1630
0
30-900
4-c.90

+
+
+

+
+

'+"

+
+
+
+
+
+

+
+

+
+

0-cA5

+
+
+
+
+

+
+
+
+
+

c.780-1730
0-21
0-c.130
0-c.90
0-18

+
+

16-88
10-1788
c.1040-2140
129-1420

Order HOLASTEROIDA
Pourtalesiidae
Stereopneustes relictus de Meijere, 1902
Urechinidae
Sternopatagus sinensis Bather, 1934

250-907

HOLOTHUROIDEA
Order ASPIDOCHlROTIDA
Holothuriidae
Actinopyga echinites (Jaeger, 1833)
Actinopyga flammea Cherbonnier, 1979
Actinopyga lecanora (Jaeger, 1833)
Actinopyga mauritiana (Quoy & Gaimard, 1833)
Actinopyga miliaris (Quoy & Gaimard, 1833)
Actinopyga obesa (Selenka, 1867)
Bohadschia argus (Jaeger, 1833)
Bohadschia marmorata (Jaeger, 1833)
Bohadschia paradoxa (Selenka, 1867)17
Bohadschia similis (Semper, 1868)18
Bohadschia tenuissima (Semper, 1868)18
Bohadschia vitiensis (Semper, 1868)18
Holothuria
(Acanthotrapeza) coluber Semper, 1868
(Acanthotrapeza) kubaryi Ludwig, 1875
(Cystipus) inhabilis Selenka, 1867
(Cystipus) dura Cherbonnier & Feral, 1981
(Cystipus) rigida (Selenka, 1867)
(Halodeima) atra Jaeger, 1833
(Halodeima) edulis Lesson, 1830
(Halodeima) pulla Selenka, 1867
(Lessonothuria) lineata Ludwig, 1875 19

+
+
+
+
+
+
+
+

+
+
+
+

+
+
+
+
+
+
+
+
+
+
+
+

'++"
+
+
+

+
+

+
+
+

+
+
+

+
+
+
+

+
+
+

+
+
+

+
+
+
+
+
+
+

+
+
+
+
+
+
+
+
+
+

+
+
+
+

+
+
+
+
+

0-5
35-60
0-23
0-12
0-20
0-25
0-40
0-36
0-41
1-16
8-25
1-15
0-25
<20
0-200
100-210
0-20
0-30
0-45
<20
0-?1O

17 occurrence beyond Hawaiian islands requires confirmation according to Rowe & Gates, 1995: 289.
18 validity of these spp. debatable; possible synonymy with Bohadschia marmorata (Jaeger, 1833) requires reexamination - see Rowe & Gates, 1995: 289.
19 The identifications in Liao & Clark, 1995 and Liao, 1997 of Holothuria (Lessonothuria) pardalis Selenka,
1867 and Holothuria (Lessonothuria) insignis Ludwig, 1875 are confused; Holothuria (L.) insignis is a junior
synonym of Hololharia (L.) pardalis, while Hololharia (L.) pardalis in Liao & Clark, 1995:438 and Liao,
1997:105 is referable to Hololhuria (Lessonothuria) lineala Ludwig, 1875 (F.W.E.R.).

488

THE RAFFLES BULLETIN OF ZOOLOGY 2000 Supplement No. 8

2
(Lessonothuria) multipilula Liao, 1975
(Lessonothuria) pardalis Se1enka, 1867 19
(Lessonothuria) verrucosa Se1enka, 1867
(Mertensiothuria) leucospilota (Brandt, 1835)
(Metriatyla) albiventer Semper, 1868
(Metriatyla) martensi Semper, 1868
(Metriatyla) scabra Jaeger, 1833
+
(Microthele) juscopunctata Jaeger, 1833
(Microthele) nobilis (Selenka, 1867)20
(Microthele) whitmaei Bell, 188720
(Platyperona) difficilis Semper, 1868
(Platyperona) samoana Ludwig, 1875
(Selenkothuria) erinacea Semper, 1868
(Selenkothuria) moebii Ludwig, 1883
(Selenkothuria) sinica Liao, 1980
(Semperothuria) cinerascens (Brandt, 1835)
(Semperothuria) flavomaculata Semper, 1868
(Semperothuria) imitans Ludwig, 1875
(Stauropora) discrepans Semper, 1868
(Stauropora) juscocinerea Jaeger, 1833
(Stauropora) olivacea Ludwig, 1888
(Stauropora) pervicax Se1enka, 1867 21
(Theelothuria) asperita
Cherbonnier & Feral, 1981
(Theelothuria) foresti
Cherbonnier & Feral, 1981
(Theelothuria) klunzingeri Lampert, 1885
(Theelothuria) notabilis Ludwig, 1875
(Theelothuria) ocellata (Jaeger, 1833)
+
(Theelothuria) spinijera Thee1, 1886
+
(Theelothuria) squamijera Semper, 1868
(Thymiosycia) arenicola Semper, 1868
(Thymiosycia) gracilis Semper, 1868
(Thymiosycia) hilla Lesson, 1830
(Thymiosycia) impatiens (Forska1, 1775)
(Vaneyothuria) integra Koehler & Vaney, 1908 .
Labidodemas pertinax (Ludwig, 1875)
Labidodemas rugosum (Ludwig, 1875)
Labidodemas semperianum Selenka, 1867
Pearsonothuria graeffei (Semper, 1868)
Stichopodidae
Stichopus chloronotus Brandt, 1835
Stichopus flaccus Liao, 1980
Stichopus hermanni Semper, 186822
Stichopus horrens Selenka, 1867
Stichopus naso Semper, 186822
Stichopus pseudohorrens Cherbonnier, 1967
Thelenota ananas (Jaeger, 1833)
Thelenota anax H.L. Clark, 1921
Thelenota rubralineata Massin & Lane, 1991

Depth range m.

Distribution

Taxa
3

+
+
+
+
+
+
+

+
+
*
+
+
+

+
+
+

+
+
+
+

+
+
+
+
+

+
+
+
+
+
+
+
+
+

+
+
+
+

+
+

O-?1O
0-30
0-10
0-36
0-100
1-25
3-12
shallow water
0-40
0-20
0-25
0-5
shallow water

+
+

0-1.5
0-40

+
+
+
+

0-40
0-10
0-20

+
+
+
+

+
+
+
+
+
+
+

174-223

122-185
intertidal
0-5
9-78
0-60
10-102
0-30
shallow water
0-220
0-30
217-230
inshore
<20
<30
5-30

+
+
+
*

+
+

+
+
+
+

+
+

+
+

+
+

+
+
+

+
+
+

+
+
+

+
+
+

+
+
+
+
+
+

+
+
+
+

+
+

+
+
+

+
+

+
+

+
+
+
+
+
+
+
+

+
+
+
+
+

0-15
28-48
0-20
0-10
0-25
8-60
0-50
1-30
14-40

20 the previously confused taxonomy of these two beche de mer species has been sorted out by Rowe & Gates,
1995: 295-296.
21 Liao & Clark' s record (1995: 443) may represent Holothuria (Stauropoda) dojleinii Augustin, 1908 (F.W.E.R.).
22 Rowe & Gates, 1995: 324-326 recognise the validity of these spp. previously included under the synonymy
of Stichopus variegatus.

489

Lane et al.: Echinoderm fauna of the South China Sea


Taxa

Depth range m.

Distribution
2

Synallactidae
Bathyplotes imperfectus
Cherbonnier & Feral, 1981
Bathyplotes punctatus (Sluiter, 1901)
Bathyplotes sulcatus Sluiter, 1901
Bathyplotes sp.
Meseres peripatus Sluiter, 1901
Meseres trachus (Sluiter, 1901)
Mesothuria crebrapedes
Cherbonnier & Feral, 1981
Mesothuria parva (Thee!, 1886)
Mesothuria regularia Heding, 1940
Mesothuria verrilli (Theel, 1886)
Paelopatides quadridens Heding, 1940
Synallactes angustus Cherbonnier & Feral, 1981
Synallactes crebrapapilla
Cherbonnier & Feral, 1981

+
+
+
+
+
+

415-510
310-614
90-1800
deep water
794-1850
180-1420

+
+
+
+

...

415-510
270-774
448-818
173-2520
750-925
415-510

415-510

OroerDENDROCHmOITDA
Psolidae
Psolus boholensis Semper, 1868
Psolus complanatus Semper, 1868
Cucumariidae
Actinocumis typicus Ludwig, 1875
Cercodemas anceps (Selenka, 1867)
Colochirus crassus Ekman, 1918
Colochirus quadrangularis Troschel, 1846
+
Colochirus robustus Oestergren, 1898
Cucumaria japonica Semper, 1868
Leptopentacta imbricata (Semper, 1868)
Leptopentacta punctabipeda Cherbonnier, 1960
Leptopentacta pygmaea (Semper, 1868)
Mensamaria intercedens (Lampert, 1885)
Plesiocolochirus armatus (van Maranzeller, 1881)23
Plesiocolochirus inornatus (van Maranzeller, 1881)
Pseudocnus echinatus (van Marenzeller, 1881)
Pseudocolochirus violaceus (Theel, 1886)
Sc1erodactylidae
Afrocucumis africana (Semper, 1868)
Cladolabes aciculus (Semper, 1868)
Cladolabes crassus (H.L. Clark, 1938)
Cladolabes schmeltzi (Ludwig, 1875)
Euthyonidiella tungshanensis (Yang, 1937)
Ohshimella ehrenbergi (Selenka, 1867)
Ohshimella nhatrangensis
Levin & Dao Tan Ho, 1989
Sclerodactyla multipes (Thee1, 1886)
Phyllophoridae
Havelockia novacorona (Cherbonnier, 1960)
Havelockia versicolor (Semper, 1868)
Havelockia spY
Hemithyone semperi (Bell, 1884)
Neothyonidium inflatum (Sluiter, 1901)

+
+
+
+
+
+
+

...

...

+
+

+
+
+

+
+
+
+

+
+
+

+
+
+
+
+
+
+

+
+

...

0-50
0-50
0-58
0-115
7-112
6-54
<20
<20
0-183
0-37
86-315
0-385
0-67
intertidal
3-8
<20
0-23
0-80
3-5

...
50

+
+
+
+

+
+
+

0-60
0-13
37-85

23 considered by F.W.E.R. to include the taxon Pentacta [ =Plesiocolochirusl nipponensis (H.L. Clark, 1938).
24 represents a number of nominal species - see Liao & Clark, 1995: 486-487.

490

THE RAFFLES BULLETIN OF ZOOLOGY 2000 Supplement No. 8

Neothyonidium magnum (Ludwig, 1882)


Phyllophorus roseus Cherbonnier & Feral, 1981
Phyllophorus (Phyllophorella) dubius
Cherbonnier, 1960
Phyllophorus (Phyllophorella) kohkutiensis
Heding & Panning, 1954
Phyllophorus (Phyllophorella) liuwutiensis
Yang, 1937
Phyllophorus (Phyllophorella) spiculata
Chang, 1935
Phyllophorus (Phyllothuria) eebuensis
(Semper, 1868)
Phyllophorus (Phyllothuria) hypsipyrga
(van Marenzeller, 1881)
Phyllophorus (Phyllothuria) ordinata Chang, 1935
Phyrella fragilis (Ohshima, 1912)
Semperiella tenera (Ludwig, 1875)
Stolus albescens Liao, 1995
Stolus bueeaUs (Stimpson, 1855)
Stolus eaneseens (Semper, 1868)
Stolus conjugens (Semper, 1868)
Thorsonia adversaria (Semper, 1868)
Thyone anomala Oestergren, 1898
Thyone bieornis Ohshima, 1915
Thyone fusus chinensis Yang, 193725
Thyone papuensis Theel, 1886
Thyone pedata Semper, 1868
Thyone pro/usus Cherbonnier & Feral, 1981
Thyone spinijera Liao, 1995
Thyone ef villosa Semper, 1868
Placothuriidae
Placothuria molpadioides (Semper, 1868)26

Depth range m.

Distribution

Taxa
3

.
...

...

0-30
70-76

<20

19-150

...

intertidal

0-30
7-125

+
+
+
+
+

...
+

...

+
+
+

+
+
+

...
+
...

+
+

<20-100
<20
<20
<20
16-109
0-54
21-89
2-147
16-50
0-103
23-61

+
+

0-60
c.55
592-610
15-115
43-73

14-57

...

...

Order DACTYLOCHIROTIDA
Ypsilothuriidae
Stauroeueumis noeturna (Sluiter, 1901)
Ypsilothuria bitentaeulata (Ludwig, 1893)

+
+

Vaneyellidae
Mitsukuriella squamulosa (Mitsukuri, 1912)27

685-757
135-4000
65-190

Order MOLPADIIDA
Caudinidae
Acaudina bacilla Cherbonnier & Feral, 1981
Acaudina leueoproeta (H.L. Clark, 1938)
Acaudina molpadioides (Semper, 1868)
Acaudina suspecta Cherbonnier & Feral, 1981
Caudina atacta Pawson & Liao, 1992
Caudina intermedia Liao & Pawson,1993
Paraeaudina ehilensis (Muller, 1850)
Paracaudina delicata Pawson & Liao, 1992

...
+
+

...

...

+
+

+
+
+

+
+

186-187
0-122
0-330
165-287
63-91
107
0-990
10-21

25 validity of this taxon awaits re-examination of type material -see Liao & C1ark, 1995: 504.
26 moved to genus Placothuria [Placothuriidae] by Liao, 1997: 224.
27 transferred from Phyllophoridae to Vaneyellidae - see Liao, 1997: 229.

491

Lane et al.: Echinoderm fauna of the South China Sea


Taxa

Distribution
2

Molpadiidae
Molpadia changi Pawson & Liao, 1992
Molpadia guangdongensis Pawson & Liao, 1992
Molpadia lenticulum (Cherbonnier & Feral, 1981)
Molpadia musculus Risso, 1826
Molpadia parvulum (Cherbonnier & Feral, 1981)
Molpadia roretzi (von Marenzeller, 1877)

+
+

Depth range m.
6

+
+

...
+

35-90
89-200
750-925
592-2340
182-510
44-620

Order APODIDA
Synaptidae
Anapta gracilis Semper, 1868
Euapta godeffroyi (Semper, 1868)
Oestergrenia dubia Semper, 1868
Oestergrenia incerta (Ludwig, 1875)
Oestergrenia variabilis Theel, 1886
Opheodesoma australiensis Heding, 1931
+
Opheodesoma clarki Heding, 1928
Opheodesoma glabra (Semper, 1868)
Opheodesoma grisea (Semper, 1868)
Opheodesoma lineata Heding, 1928
Opheodesoma spectabilis Fisher, 1907
Patinapta laevis (Bedford, 1899)
Patinapta ooplax (von Marenzeller, 1881)
Patinapta taiwanensis Chao, Rowe & Chang, 1988
Polyplectana galatheae Heding, 192828
Polyplectana grisea (Heding, 1931)
Polyplectana kefersteini (Selenka, 1867)
Polyplectana nigra (Semper, 1868)
Polyplectana samoae Heding, 193 j28
Polyplectana unispicula Heding, 1931 28
Polyplectana zamboangae Heding, 192828
Protankyra asymmetrica (Ludwig, 1875)
Protankyra bidendata (Woodward & Barrel!, 1858)
Protankyra magnihamula Heding, 1928
Protankyra pseudodigitata (Semper, 1868)
Protankyra rodea (Sluiter, 1888)
Protankyra similis (Semper, 1868)
Protankyra suensoni Heding, 1928
Protankyra verrilli (Theel, 1886)
Synapta maculata (Chamisso & Eysenhardt, 1821)
Synaptula indivisa (Semper, 1868)
Synaptula lactea (Sluiter, 1888)
Synaptula lamperti Heding, 1928
Synaptula madreporica Heding, 1928
Synaptula media Cherbonnier & Feral, 1984
Synaptula recta (Semper, 1868)
Synaptula reticulata (Semper, 1868)
Synaptula virgata (Sluiter, 1901)

...

Chiridotidae
Chiridota intermedia Bedford, 1898 29
Chiridota liberata Sluiter, 1887
Chiridota rigida Semper, 1868
Chiridota stuhlmanni Lampert, 1896
Myriotrochus sp.

+
+

...
+

+
+
+

+
+

+
+
+

+
+

+
+

+
+

...
+
+
+
+
+
+
+
+

+
+
+
+

+
+
+

...
+

+
+

...

+
+
+

+
+

+
+
+

+
+

+
+

+
+

+
+
+
+
+

<20
<20
0-10
0-5

+
+
+

28 the identification / validity of these spp. is questionable (F.W.E.R.).


29 possible synonym of Chiridota rigida Semper, 1868 - see Thandar & Rowe, 1989: 154

492

1-12
0-77
<20
20-100
17-26
15-30
<20
0-20
0-36
10-35
<20
<20
<20
0-7
<20
<20
0-77
0-36
<20
<20
<20
5-80
0-45
0-50
12-32
18-330
<20
28-90
0-14
0-25
<20
9-40
0-35
<20
8-20
0-20
9-15
18

THE RAFFLES BULLETIN OF ZOOLOGY 2000 Supplement No. 8


Taxa

Polycheira fusca (Quay & Gaimard, 1833)

Distribution
2

Depth range m.
6

<20

724-4820
174-223
309-858
1100-1301

Order ELASIPODIDA

Deimatidae
Deima validum validum Thee!, 1879
Orphnurgus bacillus Cherbannier & Feral, 1981
Orphnurgus insignis Fisher, 1907
Orphnurgus protectus Hansen, 1975

+
+

Laetmogonidae
Laetmogone violacea Theel, 1879
Pannychia moseleyi Theel, 1882

225-1804
212-2598

+
+

Psychropotidae
Psychropotes longicauda Theel, 1882

493

1100-5173

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