THE RAFFLES BULLETIN OF ZOOLOGY 2004

THE RAFFLES BULLETIN OF ZOOLOGY 2004 Supplement No. 11: 61-66

STOMATOPOD CRUSTACEA FROM ANAMBAS AND NATUNA ISLANDS,

SOUTH CHINA SEA, INDONESIA

Shane. T. Ahyong
Department of Marine Invertebrates, Australian Museum, 6 College Street, Sydney, NSW 2010, Australia
Email: shanea@austmus.gov.au

Mohammad Kasim Moosa

Research Center for Oceanography, Indonesian Institute of Sciences, Jalan Pasir Putih 1,
Ancol Timur, Jakarta Utara , Indonesia
Email: bkasim@pacific.net.id

ABSTRACT. – The stomatopod Crustacea collected during Expedition Anambas 2002 to the Anambas and
Natuna Islands, Indonesia are reported. Twelve species in seven genera and two families are reported. All
are new records for the study area. A lectotype is selected for Carinosquilla multicarinata White, restricting
the type locality to the Philippines.

KEY WORDS. – Stomatopoda, Crustacea, Anambas, Natuna, Indonesia, taxonomy.

INTRODUCTION subtilis (Manning, 1978), included in the present report, was

inadvertently excluded by Moosa (2000). With the addition
The species presented in this paper were collected during of A. orientalis, Carinosquilla thailandensis, Clorida
Expedition Anambas 2002 carried out in the southern part of albolitura, and Q. subtilis, the total known number of
the South China Sea using the Baruna Jaya VIII, the stomatopod species in the South China Sea is increased to
Indonesian Institute of Sciences’ research vessel. The 124. The present collection comprises 12 species in seven
expedition made collection around the Anambas and Natuna genera and two families. Of the 12 species reported herein,
Islands in the southern South China Sea. The expedition was 10 were listed by Moosa (2000). No stomatopods, however,
jointly sponsored by countries surrounding the South China have been recorded from the Anambas and Natuna Islands,
Sea and involved scientists from China, Indonesia, Malaysia, so all species reported here are new records for the study
Philippines, Singapore, Chinese-Taipei, Thailand, and area. Although all species reported here are new records for
Vietnam. the region, habitats frequented by small, cryptic gonodactyloid
species, such as deep crevices in living coral or coralline rock,
The biological collections were made from various types of were not sampled. Numerous species of gonodactyloids are
habitat using different collecting methods, from hand to seine abundant in reef habitats elsewhere in Indonesia (e.g., Moosa
nets. The collections were sorted at the Raffles Museum of & Erdmann, 1994; Erdmann & Sisovann, 1999) and are
Biodiversity Research of the National University of Singapore almost certainly also present in the Anambas and Natuna
and then distributed to the specialists on each taxonomic Islands. Thus, the stomatopod fauna is undoubtedly
group. underestimated. The present study, nevertheless, serves as
an initial baseline for future studies.
The most recent summary of the South China Sea fauna listed
120 species in 52 genera and 13 families (Moosa, 2000).
Several species have since been added to the South China MATERIALS AND METHODS
Sea fauna. Moosa (1991) synonymised Carinosquilla
thailandensis Naiyanetr, 1983, with Carinosquilla carinata Measurements of specimens are in millimetres (mm).
(Serène, 1950) but Naiyanetr et al. (2000) subsequently Carapace length (CL) is measured along the dorsal midline
reinstated the former species. Clorida albolitura Ahyong & and exlcudes the rostral plate. Total length (TL) is measured
Naiyanetr, 2000 was described from the Gulf of Thailand. dorsally from the rostral apex to the apices of the submedian
Alima orientalis Manning, 1978 was recorded from the teeth. Abdominal somite and thoracic somite are abreviated
southern Gulf of Thailand (Naiyanetr et al., 2000). Quollastria AS and TS respectively. Specimens are deposited in the

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 Ahyong & Moosa: Anambas and Natuna Stomatopoda.

Zoological Reference Collection of the Raffles Museum of species in the South China Sea, mostly from Vietnam, were
Biodiversity Research, National University of Singapore reviewed by Moosa (2000). Ahyong (2001) reported a
(ZRC) and Research Center for Oceanography (RCO), specimen collected from an intertidal reef flat from a cavity
Indonesian Institute of Sciences. Synonymies are not intended of Spongodes (Porifera) and two others collected by plankton
to be complete. Original citations, primary synonyms, major net from the same locality.
works, and those dealing with the South China Sea are
included. Distribution. – Andaman Sea to Malaysia, Japan, Indonesia,
Vietnam, the Philippines, Australia, New Caledonia, and
Samoa (Ahyong, 2001).
SYSTEMATIC ACCOUNT

GONODACTYLIDAE GIESBRECHT, 1910 Gonodactylus chiragra (Fabricius, 1781)

Gonodactylellus affinis (de Man, 1902) Squilla chiragra Fabricius, 1781: 515 [type locality: restricted to
Ambon, Indonesia, 3°43’S, 128°12’E, by neotype selection
Gonodactylus chiragra var. affinis de Man, 1902: 912 [type locality: (Manning, 1981: 217)].
Ternate, Moluccas, Indonesia, 0°48’N, 127°20’E]. Gonodactylus chiragra – Moosa, 2000: 408, 419-420; Ahyong,
Gonodactylus chiragra var. confinis de Man, 1902: 912, pl. 27, fig. 2001: 67-70, fig. 34.
66 [type locality: Ternate, Moluccas, Indonesia, 0°48’N,
127°20’E]. Material examined. – 1 male (TL 49 mm), Anambas: Jemaja Island,
Gonodactylus chiragra var. segregatus Lanchester, 1903: 448, pl. northern edge of Tiru Bay, opposite Punisan Island, 12 Mar.2002
23, figs. 6, 7 [type locality: Minikoi, Laccadive Islands, restricted (St. EA-ZJ01) (RCO); 1 female (TL 48 mm), Anambas: Matak
by lectotype designation (Ahyong, 2001)]. Island, south-eastern coast of the Niulwan Peninsular, off Peninting
Gonodactylellus affinis – Moosa, 2000: 407, 417-418; Ahyong, Strait, 14 Mar.2002 (St. EA JL04) (ZRC 2003.0503); 1 female (TL
2001: 46-48, fig. 21. 17 mm), Natuna: north-eastern coast of Panjang Island (north-west
of Bunguran Island), 17 Mar.2002 (St. EA JL07) (ZRC 2003.0504).
Material examined. – 1 female (TL 11 mm), Anambas: east coast
of Jemaja Island, mouth of Teluk Jebung, trawled, 13 Mar.2002 Remarks. – Gonodactylus chiragra inhabits shallow water,
(St. EA-TT04) (ZRC 2003.0501). where it forages for prey among coral boulders, especially in
the upper intertidal zone. Records of this species in the South
Remarks. – The species inhabits coarse substrates comprised China Sea are given by Manning (1995) and Moosa (2000),
of sponge, coral rubble, lithothamnion nodules, shelly grit while records for Australia are given by Ahyong (2001).
and gray sand at depths of 25-57 m as reported by Moosa
(1973). Moosa & Erdmann (1994) collected the species from Distribution. – Western Indian Ocean to Australia, Indonesia,
rubble on barren sand flats and Moosa (1991) reported New the South China Sea and French Polynesia (Ahyong, 2001).
Caledonian material from 13-80 m on moderately rough
substrates composed of fine white sand, coarse muddy sand,
coarse sand blocks and fragments with algae or Foraminifera. SQUILLIDAE LATREILLE, 1802
Ahyong (2001) reported the species from rocky and coral
reefs, crevices in rock, coral, sponge, or coralline algae from Areosquilla indica (Hansen, 1926)
the reef flat to 60-80 m depth. The distribution of G. affinis
in the South China Sea was given by Moosa (2000). Squilla indica Hansen, 1926: 12-14, pl. 1, fig. 4a-c [type locality:
Lohio Bay, Buton Strait, South East Sulawesi, Indonesia].
Distribution. – Western Indian Ocean to Australia, Indonesia, Areosquilla indica – Manning, 1976: 5-6, fig. 2; Moosa, 2000: 436.
the South China Sea and New Caledonia (Moosa, 2000;
Ahyong, 2001). Material examined. – 1 male (TL 28 mm), Natuna: west coast of
Bunguran Island, trawled, 18 Mar.2002 (St. EA TT08) (ZRC
2003.0505).

Gonodactylellus viridis (Serène, 1954)

Remarks. – The specimen of this seldom-reported species
agrees well with the lectotype in the Zoological Museum,
Gonodactylus chiragra var. viridis Serène, 1954: 6, 7, 10, 74-76,
87, fig. 13-3 [type locality: Cauda Bay, Vietnam]. University of Copenhagen, as well as the New Caledonian
Gonodactylinus viridis – Moosa, 2000: 407, 419. specimen in the Muséum National d’Histoire Naturelle, Paris.
Gonodactylellus viridis – Ahyong, 2001: 63-65, fig. 31.
Distribution. – Maldives, Andaman Sea, Indonesia, and New
Material examined. – 4 males (TL 26-35 mm), Anambas: Jemaja Caledonia (Moosa, 1991). The presence of this species from
Island, northern edge of Tiru Bay, opposite Punisan Island, 12
the South China Sea was questionably reported by Moosa
Mar.2002 (St. EA-ZJ01) (3 males to ZRC 2003.0502, 1 male to
RCO). (2000) based on the report of Nguyen & Pham (1995). The
present specimen of A. indica constitutes a new record for
Remarks. – Gonodactylellus viridis is a shallow water the study area and confirms its presence in South China Sea
species, commonly found on the reef flat. The records of this waters.

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Carinosquilla carinata (Serène, 1950) upturned, straight to sinuous; apex truncate to rounded; with
long, distinct, median carina flanked by long supplementary
Squilla carinata Serène, 1950: 571 [type locality: Cauda Bay, carina. Carapace anterior width 0.52 CL; anterior bifurcation
Nhatrang, Vietnam]. of median carina opening posterior to dorsal pit. Raptorial
Carinosquilla carinata – Naiyanetr, 1980: 43; Manning, 1995: 173, claw dactylus with 5 teeth; merus outer face with longitudinal
figs. 101-104, 105b, 106b, 107b, 108b, pl. 30; Moosa, 2000: carina. Mandibular palp present. Maxilliped 1-4 with epipod.
436.
TS5 lateral process with anterior lobe a slender spine directed
Material examined. – 2 males (TL 86-89 mm), 2 females (TL 93- anterolaterally; posterior lobe short, broad with rounded apex
97 mm), Anambas: Tarempa Bay, 14 Mar.2002 (St. EA TT-06) directed laterally. TS6 lateral process anterior lobe quadrate,
(ZRC 2003.0506).

Remarks. – The present specimens agree well with published

accounts and exhibit typical variation in the armature of the
raptorial claw. The largest female bears six teeth on the
dactylus of one raptorial claw and seven on the other. All
other specimens have six teeth on the dactyli of both claws.

Carinosquilla carinata inhabits mud or muddy-sand

substrates between 15 and 50 m depth. Moosa’s (1991)
records of C. carinata from New Caledonia are based on C.
australiensis Ahyong, 2001, and C. redacta Ahyong, 2001.

Distribution. – Vietnam, the Gulf of Thailand, and now from

Indonesia. The distribution of this species in Vietnam and
Thailand is given by Moosa (2000).

Carinosquilla multicarinata (White, 1849)

(Fig. 1)

Squilla multicarinata White, 1849: 144, pl. 6, fig. 1 [type locality:

the Philippines, fixed by present lectotype designation].
Carinosquilla multicarinata – Naiyanetr, 1980: 43; Moosa, 2000:
437.

Material examined. – 1 male (TL 68 mm), Anambas: Tarempa Bay,

14 Mar.2002 (St. EA TT-06) (RCO); 1 male (TL 59 mm), 1 female
(TL 81 mm), Anambas: Matak Island, south-eastern coast of the
Niulwan Peninsular, off Peninting Strait, 14 Mar.2002 (EA TT04)
(ZRC 2003.0507).

Diagnosis. – Eyestalk without carinae. Ocular scales entire,

not bifurcate. Carapace anterior with anterior bifurcation of
median carina opening posterior to dorsal pit. Raptorial claw
dactylus with 5 teeth; merus outer face with longitudinal
carina. Mandibular palp present. TS5 dorsal carinae
transverse, except medially. TS6-8 and AS1-6 dorsal carinae
subparallel, most or all posteriorly armed above intermediate
carinae. AS1-6 with supplementary carinae unarmed below
intermediate carinae. Abdominal somites with normal
complement of carinae spined as follows: submedian 1-6,
intermediate 1-6, lateral 1-6, marginal 1-5. Telson prelateral
lobe with sharp apex; dorsolateral surface with numerous
supplementary longitudinal carinae, uninterrupted proximally.
Uropodal protopod inner margin with slender spines.
Uropodal exopod exopod distal segment black.

Description of lectotype. – Eyestalk without carinae. Ocular

scales entire, not bifurcate. A1 somite dorsal processes with Fig. 1. Carinosquilla multicarinata (White, 1849). Lectotype,
acute apices; directed anterolaterally. A2 scale length 0.55 female, TL 67 mm, BMNH 709, Philippine Islands (photo: H.
CL. Rostral plate as long as broad; lateral margins convergent, Taylor).

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apex rounded to subtruncate; posterior lobe broad, rounded. Distribution. – Gulf of Thailand, northeastern Australia, and
TS7 lateral process anterior lobe short, blunt; posterior lobe now from Anambas, Indonesia.
broad, rounded, larger than anterior lobe. TS8 anterolateral
margin triangular, apex acute; sternal keel triangular. TS5
dorsal carinae transverse, except medially. TS6-8 and AS1- Cloridina chlorida (Brooks, 1886)
6 dorsal carinae subparallel, most or all posteriorly armed
above intermediate carinae. AS1-6 with supplementary Squilla chlorida Brooks, 1886: 21, 40, pl. 2, figs. 1-5 [type locality:
carinae unarmed below intermediate carinae. AS6 submedian Amboina, Indonesia, 3°43’S, 128°12’E, 27 m].
carinae unicarinate with supplementary longitudinal carinae Cloridina chlorida – Moosa, 2000: 439; Ahyong, 2001: 232-233,
laterally and medially; sternum with continuous transverse fig. 113.
proximal carina and a V-shaped median carina flanked by
Material examined. – 1 male (TL 46 mm), Anambas: Teluk Tarempa
transverse carina. Abdominal somites with normal
Bay, 03°15.31-15.28’N, 106°09.50-11.79’E, trawled, 46 m, 14
complement of carinae spined as follows: submedian 1-6,
Mar.2002 (St. EA-TT-06.) (ZRC 2003.0509).
intermediate 1-6, lateral 1-6, marginal 1-5. Telson as long as
broad; prelateral lobe longer than margin of lateral tooth, with
Remarks. – The single specimen agrees well with published
sharp apex; dorsolateral surface with numerous accounts (Brooks, 1886; Manning, 1968; Ahyong, 2001).
supplementary longitudinal carinae, uninterrupted proximally; Cloridina chlorida occupies muddy substrates at depths of
denticles submedian 3, intermediate 8-9, lateral 1. Uropodal 30-64 m (Manning, 1968).
protopod inner margin with 8 slender spines; with ventral
tubercle anterior to endopod articulation; protopod terminal Distribution. – Madagascar, Vietnam, Indonesia, the
spines with lobe on outer margin of inner spine rounded, as Philippines, New Caledonia and Australia (Ahyong, 2001).
broad as or narrower than adjacent spine, proximal margin
concave. Uropodal exopod proximal segment outer margin
with 9 movable spines, distalmost not exceeding midlength Oratosquillina perpensa (Kemp, 1911)
of distal segment, distal margin with 2 slender ventral spines
outer longest; exopod distal segment black, ventrally carinate, Squilla oratoria var. perpensa Kemp, 1911: 98 [part][type locality:
length subequal to proximal segment; endopod dorsally and Hong Kong, restricted by lectotype designation (Manning,
ventrally carinate. 1978)].
Squilla oratoria var. inornata. – Holthuis, 1941: 248 [part, not
Remarks. – We take this opportunity to designate the female Squilla oratoria var. inornata Tate, 1883].
syntype from the Philippines as the lectotype. The lectotype Oratosquilla perpensa – Manning, 1978: 21-23.
is a 67 mm TL dry female specimen in the collections of the Oratosquillina perpensa – Manning, 1995: 233-234; Moosa, 2000:
Natural History Museum, London, registration number 709. 449.
The increasing numbers of species recognized in
Carinosquilla, including a species nearly identical to C. Material examined. – 2 females (TL 82-102 mm), Natuna: west
multicarinata (see Ahyong, 2001), and that C. multicarinata coast of Bunguran Island, 18 Mar.2002 (St. EA TT08) (1 female to
is the type species of the genus justifies the present action. ZRC 2003.0510, 1 female to RCO).

Distribution. – Eastern Indian Ocean to Vietnam, the Gulf Remarks. – The specimens agree well with Manning’s (1978)
of Thailand, Indonesia, the Philippines and Japan. Moosa redescription. Although variation in the length of the rostral
plate has been observed in O. perpensa (e.g., Ahyong, 2001;
(2000) presented records of this species in the South China
Ahyong & Naiyanetr, 2002) both specimens examined here
Sea.
have the ‘short’ rostral plate of the lectotype.

Distribution. – Burma to Indonesia, Vietnam, Hong Kong,

Carinosquilla thailandensis Naiyanetr, 1983 Taiwan and Japan (Manning, 1995). The distribution within
the South China Sea was given by Moosa (2000).
Carinosquilla thailandensis Naiyanetr, 1983: 394-399, figs. 2, 4
[type locality: Ko Phai, Chon Buri province, Thailand, 12°56’N,
100°41’E]; Naiyanetr et al., 2000: 1292-1294; Ahyong, 2001: Oratosquillina quinquedentata (Brooks, 1886)
214, fig. 104.
Squilla quinquedentata Brooks, 1886: 21, 26, pl. 1: fig. 3, pl. 2: fig.
Material examined. – 2 males (TL 104-105 mm), 1 female (TL 81 6 [type locality: Arafura Sea, 09°59’S, 139°42’E].
mm), Anambas: east coast of Jemaja Island, mouth of Teluk Jebung, Oratosquillina quinquedentata – Moosa, 2000: 449; Ahyong, 2001:
13 Mar.2002 (St.EA TT04) (1 male, 1 female to ZRC 2003.0508, 295-298, fig. 144.
1 male to RCO).
Material examined. – 1 male (TL 112 mm), Anambas: east coast
Remarks. – The three specimens agree well with published of Jemaja Island, mouth of Jebung Bay, 13 Mar.2002 (St. EA TT04)
accounts (Naiyanetr, 1983; Naiyanetr et al., 2000; Ahyong, (ZRC 2003.0511).
2001) and constitute the first records of the C. thailandensis
from Indonesia. The species was previously known only from Remarks. – Oratosquillina quinquedentata burrows in muddy
the Gulf of Thailand and northeastern Australia, so the present or sandy-mud substrates from the shore to at least 51 m depth
records are the first from an intermediate locality. (Ahyong, 2001).

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Distribution. – India to the Gulf of Thailand, Indonesia, and LITERATURE CITED

Australia (Ahyong, 2001).
Ahyong, S. T., 2001. Revision of the Australian Stomatopod
Crustacea. Records of the Australian Museum, Supplement 26:
Quollastria gonypetes (Kemp, 1911) 1-326.
Ahyong, S. T. & P. Naiyanetr, 2002. Stomatopod Crustaceans from
Squilla gonypetes Kemp, 1911: 96 [type locality: vicinity of Cheduba Phuket and the Andaman Sea. Phuket Marine Biological Centre
Island., Burma, 18°48’N, 93°38’E, 13 m, restricted by lectotype Special Publication, 23(2): 281-312.
designation (Manning, 1978)]. Brooks, W. K., 1886. Report on the Stomatopoda collected by
Oratosquillina gonypetes – Moosa, 2000: 447. H.M.S. Challenger during the years 1873-76. The Voyage of
Quollastria gonypetes – Ahyong, 2001: 304-306, fig. 147. the H.M.S. Challenger, Zoology, 16: 1-116, pls. 1-16.
Erdmann, M. V. & O. Sisovann, 1999. Distribution and abundance
Material examined. – 1 female (TL 68 mm), Natuna: west coast of of reef-flat stomatopods in Teluk Jakarta and Kepulauan Seribu.
Bunguran Island, 18 Mar.2002 (St.EA TT-08) (ZRC 2003.0512). In: Soemodiharjo, S. (ed.), Proceedings of Coral Reef Evaluation
Workshop, Pulau Seribu, Indonesia, Study 10. UNESCO, pp.
Remarks. – The specimen agrees well with published 66-83.
accounts, exhibiting the distinctive pair of dark ‘squares’ on Fabricius, J. C., 1781. Species Insectorum Exhibentes Eorum
the fifth abdominal somite. This species inhabits muddy sand Differentias Specificas, Synonyma Auctorum, Loca Natalia,
with Foraminifera, with depth ranging from 13 to 180 m Metamorphosin Adiectis, Observationibus, Descriptionibus 1.
(Moosa, 2000). Hamburgii et Kilonii. 552 pp.
Giesbrecht, W., 1910. Stomatopoden, Erster Theil. Fauna und Flora
Distribution. – Western Indian Ocean to Australia, Indonesia, des Golfes von Neapel Monographie 33: i-vii, 1-239, pls. 1-11.
the South China Sea, and Japan (Ahyong, 2001). Hansen, H. J., 1926. The Stomatopoda of the Siboga Expedition.
Siboga-Expeditie, monographie, 35: 1-48, pls. 1, 2.
Holthuis, L. B., 1941. The Stomatopoda of the Snellius Expedition.
Quollastria subtilis (Manning, 1978) Biological Results of the Snellius Expedition XII. Temminckia,
6: 241-294.
Oratosquilla subtilis Manning, 1978: 33-34, fig. 19 [type locality:
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1991: 12.
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carried on and of the collections made by an expedition during
Material examined. – 2 females (TL 43-46 mm), Anambas: Teluk the years 1899 and 1900, 1: 444-459.
Tarempa Bay, 14 Mar.2002 (St. EA TT-06) (ZRC 2003.0513); 1
Latreille, P. A., 1802. Histoire naturelle, générale et particulière,
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Mar.2002 (St. EA TT-08) (RCO).
Man, J. G. de., 1902. Die von Herrn Professor Kükenthal im
Indischen Archipel gesammelten Dekapoden und Stomatopoden.
Remarks. – The specimen agrees well with published In: Kükenthal, W. (ed.), Ergebnisse einer zoologischen
accounts, exhibiting the characteristic pair of dark ‘triangles’ Forschungsreise in den Molukken und Borneo. Abhandlungen
on the fifth abdominal somite. Quollastria subtilis occupies der Senckenbergischen naturforschenden Gesellschaft, 25: 467-
sand or silty substrates and is known from depths of 31-111 929, pls. 19-27.
m (Ahyong, 2001). Manning, R. B., 1968. Stomatopod Crustacea from Madagascar.
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Distribution. – Western Indian Ocean to Australia, Indonesia,
Manning, R. B., 1976. Redescriptions of Oratosquilla indica
the South China Sea, and New Caledonia (Ahyong, 2001).
(Hansen), with accounts of a new genus and two new species
(Crustacea, Stomatopoda). Beaufortia, 25(318): 1-13.
Manning, R. B., 1978. Further observations on Oratosquilla, with
ACKNOWLEDGMENTS
accounts of two new genera and nine new species (Crustacea:
Stomatopoda: Squillidae). Smithsonian Contributions to
We are grateful to Peter Ng (National University of Zoology, 272: 1-44.
Singapore) for making the collections available for study, and
for providing us with financial support and working space at Manning, R. B., 1981. Neotype selection for the stomatopod Squilla
chiragra Fabricius, 1781. Crustaceana, 40 (4): 217-219.
the Raffles Museum of Biodiversity Research during the
completion of this study. Miranda Lowe and Harold Taylor, Manning, R. B., 1995. Stomatopod Crustacea of Vietnam: the legacy
both of the Natural History Museum, London, are gratefully of Raoul Serène. Crustacean Research, Special No. 4: 1-339.
acknowledged for excellent photographs of the lectotype of The Carcinological Society of Japan. Shimoda Printing,
Kumamoto, Japan.
C. multicarinata.

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