THE RAFFLES BULLETIN OF ZOOLOGY 2004
THE RAFFLES BULLETIN OF ZOOLOGY 2004 Supplement No. 11: 79-88

BRACHYURA (CRUSTACEA) OF THE ANAMBAS EXPEDITION 2002
Darren C. J. Yeo
Raffles Museum of Biodiversity Research, Department of Biological Sciences, National University of Singapore,
10 Kent Ridge Crescent, Singapore 119260
Email: darrenyeo@nus.edu.sg

Dwi Listyo Rahayu
Research Centre for Oceanography, Indonesian Institute of Sciences (LIPI), Jalan Pasir Putih 1,
Ancol Timur, Jakarta 11048, Indonesia
Email: dwilistyo@yahoo.com

Peter K. L. Ng
Raffles Museum of Biodiversity Research, Department of Biological Sciences, National University of Singapore,
10 Kent Ridge Crescent, Singapore 119260
Email: peterng@nus.edu.sg

ABSTRACT. – The brachyuran crabs collected by the 2002 scientific expedition to the Anambas and Natunaislands in the South China Sea are reported upon. A total of 87 species from 15 families are recorded.
Noteworthy are the discovery of two new species, one pilumnid, Heteropilumnus satriai, and one sesarmid,
Parasesarma anambas.
KEY WORDS. – Taxonomy, Anambas, South China Sea, Heteropilumnus, Parasesarma, new species, records.

treated in detail in this paper (marked in Appendix 1 by anasterisk).

INTRODUCTION
In March 2002, Expedition Anambas, was conducted in thesouthern part of the South China Sea, as part of the regional
workshop process in resolving conflicts through confidencebuilding measures. The expedition, which lasted a fortnight,
centred on the relatively poorly explored Indonesian islandsof Anambas and Natuna, and included scientific participantsfrom Indonesia, Singapore, Malaysia, Philippines, Thailand,
Vietnam, China and Chinese-Taipei. Specimens wereobtained by trawling, as well as by hand-collecting during
diving and intertidal beach-combing.

TAXONOMIC NOTESFAMILY MAJIDAE SAMOUELLE
Austrolibinia pincerna Wagner, 1992
Remarks. – The taxonomy of this characteristic species has
been discussed in depth by Wagner (1992) and Rahayu &Ng (2000). The type locality is in the South China Sea, near
the Anambas (Wagner, 1992), and the rediscovery of thespecies here is thus noteworthy. It has also been reported fromthe Indonesian part of the Straits of Malacca (Rahayu & Ng,
2000).

The present paper reports on the brachyuran crabs collected
as a result of this expedition. A total of 87 species from 15
families are documented (see Appendix 1). The material
examined is also listed in Appendix 1, and the datasets
associated with this material are presented in the Scientific
Overview chapter by Ng et al. (2004) (in the present volume).
Specimens are deposited in the Museum Zoologicum
Bogoriense (MZB), Cibinong, Indonesia; and Zoological
Reference Collection (ZRC) of the Raffles Museum of
Biodiversity Research, National University of Singapore.
Measurements provided in the text are of the carapace breadthand length respectively. The abbreviations G1 and G2 areused for the male first and second pleopods respectively. Only
the taxonomy of new species or those of specific interest are

FAMILY PORTUNIDAE RAFINESQUE
Subfamily Portuninae RafinesquePortunus (Xiphonectes) tenuipes (De Haan, 1835)
Remarks. – The present specimens from Anambas are clearlyreferable to Portunus tenuipes s. str., matching the description

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Yeo et al.: Anambas Expedition Brachyura
30.4 by 17.0 mm) (ZRC 1992.10341-10346), Thailand, off Pattaya,
coll. P. K. L. Ng & L. B. Holthuis, 1991. SINGAPORE: 4 males(largest 34.7 by 22.6 mm), 4 females (including 1 ovigerous, 31.2
by 14.7 mm) (ZRC 1965.7.5.37-46), Siglap, coll. M. W. F. Tweedie,
1934; 2 males (larger 36.2 by 22.0 mm) (ZRC 1985.896-897), Siglap,
coll. M. W. F. Tweedie, 1934; 1 juvenile (ZRC 1991.9575), Pulau
Semakau, coll. 1990; 1 juvenile (ZRC 1991.16181), Pulau Sakra,coll. 1991.

and illustrations of the holotype by De Haan (1835: Pl. 1 fig.
4) and Yamaguchi & Baba (1993: Fig. 138) very well. In
addition, the ZRC has some P. tenuipes specimens collectedfrom the Gulf of Thailand (see Comparative Material below)that agree well with the present Anambas material (see De
Haan, 1835: Pl. 1 fig. 4; Sakai, 1976: Fig. Pl. 120 fig. 3;
Yamaguchi & Baba, 1993: Fig. 138).
Stephenson & Campbell (1959) and Stephenson (1972)
describe P. tenuipes as possessing only a single spine on the
carpus (on the inner margin) of the cheliped; and theiridentification keys distinguish P. tenuipes from the closely
related P. rugosus by the presence of an inner spine only on
the carpus of the cheliped (versus presence of both inner andouter spine). However, the present specimens, which matchthe illustrations of P. tenuipes in Stephenson & Campbell(1959) and Stephenson (1972), do possess a spine each on
both the inner and outer margins of the carpus of the cheliped,
although the outer margin spine is relatively low. The
holotype similarly possesses a low but distinct spine on theouter margin of the carpus of the cheliped (see De Haan, 1835:
Pl. 1 fig. 4; Yamaguchi & Baba, 1993: Fig. 138). It is possiblethat the Australian specimens of P. tenuipes were describedas such by Stephenson & Campbell (1959) and Stephenson(1972) because of the orientation of the cheliped during
examination, drawing or photographing causing the alreadylow spine to almost disappear from dorsal view.
Alternatively, if the absence of the outer carpal spine is realand consistent, Stephenson & Campbell (1959) andStephenson (1972) might well be dealing with another species
altogether. A re-examination of the Australian P. tenuipes
specimens is needed to resolve this.

FAMILY PILUMNIDAE SAMOUELLE
Subfamily Rhizopinae Stimpson
Heteropilumnus satriai, new species
Material examined. – Holotype - male (19.3 by 13 mm) (MZB),
St. EA-D 11, Natuna: rocky islet southeast of Pulau Laut, 16
Mar.2002.

Description. – Carapace 1.4 times broader than long, moreor less flat across mesogastric and branchial regions; convex
fore and aft over frontal third. Carapace surface with scattered
granules on metagastric region, with denser granules onposterolateral region; scarce short setae on metagastric region,
denser anterolaterally and posterolaterally. Regions relatively
poorly defined, with only gastric region being moderately
distinct.
Front deflexed, bilobed, a median cleft with groove present;
margin sinuous with obtuse preorbital angle; postfrontal withtransverse row of minutes granules. Supraorbital margin
sinuous, granular, with two fissures. Infraorbital margin
minutely granular. Ocular peduncles short, corneas small,
darkly pigmented. Antennules relatively short, folding into
oblique fossae. Antennae with basal segment just touching
front. Anterolateral margin consisting of 4 lobes including
exorbital angle with slightly raised rim. Exorbital angle small,
subacute, with small accessory granules on outer margin; 2ndlobe large, margin with several granules; third and fourthlobes about the same size, large, margin with several granules.
Greatest carapace width between 4th anterolateral lobe.
Posterolateral margin longer than anterolateral, rounded,converging posteriorly. Posterior margin straight, with slightly
raised rim.

We also examined a good series of comparative material
collected from Singapore, which had been identified as P.
tenuipes. These, however, consistently differ from the present
Anambas and Thai material as well as from illustrations of
De Haan (1835: Pl. 1 fig. 4), Stephenson (1972), Sakai (1976:
Fig. Pl. 120 fig. 3), and Yamaguchi & Baba (1993: Fig. 138),
in G1 structure, carapace sculpturing, and limb length andproportions. In addition, they appear to be generally smaller
in size. A female specimen from Singapore ([ZRC1965.7.5.37) is already ovigerous at 31.2 by 14.7 mm (versus
an immature female specimen from Thailand [ZRC1992.10341], with the abdomen not fully expanded,measuring 30.4 by 17.0 mm). The smallest male specimen
from among the Singapore material (ZRC 1965.7.5.46),
measuring 22.8 by 10.5 mm, possessed fully developed G1s
and abdomen (versus an immature male from among the Thaimaterial [ZRC 1992.10346], with incompletely developed
G1s and abdomen, measuring 26.0 by 14.0 mm). It is possiblethat the Singapore material might represent an undescribedor poorly known species, but this is outside the scope of thepresent paper, and will need to be dealt with in a separate
work.

Third maxilliped with merus 1.5 times wider than long and
0.7 times as long as ischium; distal margin straight; ischium
about 1.4 times longer than wide, inner margin straight.
Chelipeds subequal, right slightly larger. Merus short,
trihedral, with row of minutes spines on ventral anterior
margin; outer surface with few granules near upper border,
scarcely setose; upper border smooth, with long dense setae
distally, inner border with long dense setae distally. Carpus
unarmed, inner angle blunt; surface covered with large, blunt
tubercles and short setae. Palm excluding fixed finger c. 1.2
times longer than high; fingers relatively short, fixed finger
about as long as palm, down turn. Dactyl1.2 length of palm;
both fingers with tips obtuse, cutting edges with large, obtuse
teeth, distal half of fingers darkly pigmented. Entire outer

Comparative material. – Portunus tenuipes (De Haan, 1835) –THAILAND: 1 male (43.9 by 25.8 mm) (ZRC 1970.2.19.3), Gulfof Thailand, coll. Fisheries Research Laboratory, Bangkok, 1970;
1 immature male (26.0 by 14.0 mm), 4 immature females (largest

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Fig. 1. Heteropilumnus satriai, new species. a. right anterolateral margin. b. right orbital area, ventral view, c. right third maxilliped, d.
abdomen, e, f. left G1, g. right coxa of fifth walking leg. Scale 1 mm.

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Yeo et al.: Anambas Expedition Brachyura
surface covered with large, obtuse tubercles and scarce short,
stiff setae.

The only specimen available for this study is infected by a
Sacculina (Rhizocephala), and as such, its abdomen is notnormal. It possesses both male and female pleopods, and theG1 is here figured, although it may be somewhat abnormal.
The parasite, however, does not affect the other charactersused in this study.

Walking legs fringed with long setae on dorsal and ventral
margins; upper lateral half of propodi, lateral surface of carpi
and meri with scattered long and short setae. Second and third
walking legs longest, about as long as carapace width. Dactyli
styliform, about as long as propodi. Merus of third walking
leg 3.3 times longer than wide. Small projecting scale on
upper surface of coxae of all four pairs of walking legs.

FAMILY SESARMIDAE DANA
Parasesarma anambas, new species

Thoracic sternum and abdomen scarcely setose. Telson longer
than other segments. Segment 1 widest, shortest. G1 sinuous,
apical part relatively long, curved; G2 sigmoid, very short.

Material examined. – Holotype - 1 male (8.1 by 6.9 mm) (MZB),St.EA-DW 04, Anambas: Pulau Jemaja: Teluk Jebung: northernmangrove inlet, 13 Mar.2002.

Etymology. – The species is named in honour of Mr Satria
Djambek (Indonesian Department of Foreign Affairs), whose
tireless and patient efforts have helped make the expeditiona clear success.

Paratypes – 1 male (8.1 by 7.1 mm), 1 female (4.2 by 3.4 mm)(ZRC), 2 females (4.9 by 3.8 mm; 4.8 by 4.0 mm) (MZB), St. EAJL 05, Anambas, eastern Pulau Siantan: Teluk Baruk: Sungei
Temburun, waterfall (= Air Terjun), 15 Mar.2002.

Remarks. – The new species is only tentatively referred to
Heteropilumnus De Man, 1895, owing to the distinctly more
ovate carapace (rather than the usual rectangular to
subrectangular form) and the sparsely setose carapace inwhich most of the surface is visible. In all other
Heteropilumnus species, the carapace is covered with a dense
coat of long and/or short setae such that much of the carapace
surface is obscured. In many species, the setae are also much
longer and denser on the anterior part of the carapace, so muchso that the orbits and eyes are not visible from dorsal view.
The anterolateral margin of the present species is also
distinctly convex (vs. gently convex) with very low lobes
throughout. This combination of features easily distinguishes
this species from all congeners (see also Ng & Tan, 1988;
Ng & Davie, 1991; Davie & Humpherys, 1997), and is here
described as new, Heteropilumnus satriai.

Description. – Carapace 1.25 broader than long; mesogastricand cardiac regions well defined, intestinal region moderately
defined; lateral carapace surface lined with strong obliquestriae; carapace surface very finely granular, with scarce setaeon epigastric region. Postfrontal region separated into 4 lobes
by narrow grooves, median lobes larger than lateral lobes.
Front deflexed downward, margin bilobed in dorsal view,each lobe broadly convex, separated by very broad medianconcavity. Supraorbital margin gently convex, entire. External
orbital tooth triangular, directed obliquely outwards andrepresenting point of greatest width; fused with entire lateralcarapace margin; antero- and posterolateral margins notdemarcated, without trace of tooth or indentation; lateral
margin gently sinuous, subparallel along most of length before
curving to join straight posterior carapace margin. Eyes
extending slightly beyond edge of external orbital tooth.
Antennal and antennular basal segments adjacent, notseparated by septum; basal antennular segment swollen.
Antennal flagellum moderately long, entering orbit.

Heteropilumnus satriai on the other hand, bears a marked
resemblance to some species of Cryptolutea Ward, 1936 (typespecies Cryptolutea lindemanensis Ward, 1936). Especiallywith regards to the carapace shape, anterolateral margin
armature and sparsely setose carapace, H. satriai resembles
C. arafurensis Davie & Humpherys, 1997, and C. sagamiensis(Sakai, 1935). However, H. satriai does not have the dorsal
part of the ambulatory coxa prominently produced to partiallycover the basis-ischium and with the lateral margin distinctly
serrated. In H. satriai, the ambulatory coxa has the dorsal
part slightly produced outwards (more so than in other
Heteropilumnus species examined) but clearly notoverlapping the adjacent segments, and the lateral margin issmooth. Moreover, in Cryptolutea species, the enlarged coxal
plate is distinct on all the ambulatory legs, whereas in H.
satriai, the slightly enlarged coxa is only discernible on thelast leg. For this reason, we keep it in Heteropilumnus. In
any case, it has been commented before that Heteropilumnus(type species Heteropilumnus stormi De Man, 1895),
currently with 18 valid species, is heterogeneous (see Ng,
1987), and all indications are that a revision will split it into
several genera. When this is done, the generic position of H.
satriai can be re-evaluated.

Ischium of third maxilliped with shallow median sulcus,
merus with distinct submedian ridge; exopod slender, tipreaching to half length of outer margin of merus, flagellum
long. Inner margin of merus and ischium with long setae,
proximal outer margin of ischium and base of exopod withlong, dense setae.
Cheliped subequal, robust. Merus with posterior bordercarinate, minutely tuberculate, without subdistal spine;
anterior border with minute spines ending in large subdistal
spine; outer surface with oblique striation, inner surface withscarce setae. Carpus with inner angle not produced, outermargin and dorsal surfaces striated. Upper surface of palm
with 3 transverse pectinated crests; primary crest composedof 19 tall narrow teeth; secondary crest well developed,
shorter than primary, with 10, broader and more widely
spaced teeth; tertiary crest composed of 7 much lower,
broader, widely spaced teeth, followed by several blunt
tubercles. Area below crests with rows of small tubercles.
Outer surface of palm striated proximally, smooth distally,82

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Fig. 2. Parasesarma anambas, new species. a. carapace. b, c. left cheliped, d. fifth right walking leg, e. left G1. Scale 1 mm.

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Yeo et al.: Anambas Expedition Brachyura
cheliped in P. anambas compared to only two in P. lenzi.
Furthermore, in P. lenzi, the propodus of the third leg is about
1.6 times the length of the dactyl, while in P. anambas, thethird dactyl is as long as the propodus (compared to specimensof P. lenzi in the ZRC from Singapore and Thailand).

naked; inner surface of palm with several tubercles. Fixedfinger rounded, smooth on outer surface; without ventral
ridge, moderately long. Cutting edge of fixed finger and dactylwith small teeth, larger teeth medially and distally. Dorsal
surface of dactyl with 17 symmetrical tubercles, all distinct,
small, moderately spaced proximally, becoming larger andmore widely spaced distally. Several low tubercles on
proximal third of inner edge of dorsal surface. Fingers withtips chitinous; males with narrow gape when fingers closed.

FAMILY PLAGUSIIDAE DANA
Plagusia squamosa (Herbst, 1790)

Walking legs long, slender, flattened; second and third pairs
subequal, longer than others, about 1.7 times carapace width.
Merus of third leg 2.4 times as long as wide; anterior marginof merus with acute subdistal spine. Meri of legs 1-4 withtransverse striae on upper surfaces. Carpi of legs 1-4 with 2
accessory carinae on outer surface. Propodus of third leg about
3 times as long as wide with accessory carina on inferior
proximal portion of outer surface, dorsal and ventral margins
with prominent long stiff setae. Dactyl of third leg about samelength as propodus, slightly recurved, terminating in acutecalcareous tip, dorsal and ventral margins with prominent long
stiff setae.

Remarks. – This is a well known and widely distributed IndoWest Pacific species previously known as P. tuberculata
Lamarck, 1818. Schubart & Ng (2000) recently demonstrated
that P. squamosa (Herbst, 1790) is an older name.

FAMILY GECARCINIDAE MACLEAY
Cardisoma carnifex (Herbst, 1796)
Ecological note. – Despite its occurrence in mangroves,
Cardisoma carnifex was thought to have strongly terrestrial
habits, preferring to avoid being submerged in water.
However, specimens in the present study were collected fromboth dry as well as inundated parts of a small patch of sandy
mangrove, adjacent to a small river mouth. Collection was
carried out at night during a rapidly rising tide, and during
this time, numerous individuals in the local population of C.
carnifex were observed to be submerged in water up to 50
cm deep, foraging and sheltering amongst debris, apparently
without inhibition (DCJY, pers. observ.).

Surface of thoracic sternite scarcely setose. Sternites 1-3
completely fused. Sternites 3 and 4 separated by low ridge
lined with short, scarce, setae. Abdominal cavity reaching
just below low ridge which separates sternites 3 and 4.
Male abdomen relatively broad. Telson semicircular, evenly
rounded, as long as preceding segment; segment 6 about 0.4
times as long as wide, lateral margins slightly convex.
Segments 3-5 progressively more trapezoidal, lateral margins
of segment 3 and 6 slightly convex, lateral margins ofsegments 4 and 5 straight, segments 1 and 2 very narrow
longitudinally.

GENERAL DISCUSSION
While the number of species collected by the expedition is
not very substantial, it is still interesting as two new species
were discovered. Most of the species reported here have wide
geographic distributions in the Indo-West Pacific and theirpresence in the Anambas and Natuna islands is not surprising.
In any case, there are few historical records of decapod
crustaceans from these islands and their nearby areas. Bott
(1970) described an endemic freshwater crab from Pulau
Natuna Besar (formerly Pulau Bunguran or Bunguran Island),
Balssiathelphusa natunaensis Bott, 1970 (Parathelphusidae),while Ng (1986) reported a new species, Geosesarma insulare(Sesarmidae), from Anambas; but otherwise, hardly anythingis known about the freshwater and marine fauna. We haveon hand, good material of freshwater crabs and prawns fromthese islands, and there are a good number of new genera,
new species and new records that will be reported later in a
separate paper.

G1 relatively slender, straight; apical process bent to form
an angle of 45º, strongly produced, corneous part long withtruncate tip. Setae long, simple, originating at base of apical
process and palp. G2 very short.
Female with chelipeds relatively smaller, primary pectinatedcrest on palm distinct, secondary and tertiary crests not welldeveloped, dactylar tubercles indistinct.
Etymology. – The species is named after the type locality;
used as a noun in apposition.
Remarks. – The form of the G1 of this new species resembles
Parasesarma exquisetum Dai & Song, 1986, in having the
apical process bent at an angle of 45º, with the distal corneouspart elongate with a truncate tip. However, the G1 of the latter
species is proportionately stouter and the upper margin of
the corneous part is convex, whereas in P. anambas, it is more
slender and the upper margin of the corneous part is straighter(cf. Dai & Song, 1986). In addition, P. exquistum has two
pectinated crests on the palm of the cheliped while P.
anambas has three. The form of the dactylar tubercles, which
are well spaced, also resemble those of P. lenzi De Man, 1895.
However, there are three pectinated crests on the palm of the

ACKNOWLEDGEMENTS
We are grateful to all the facilitators and participants of
Expedition Anambas, and the crew of the Baruna Jaya VIII,
for helping in the collection of specimens and data. Thanks
are also due to Ho Ping-Ho (Taiwan), Daisy Wowor, Tan
Swee Hee, Joelle Lai, Yeo Keng Loo, Leong Tzi Ming and
Lim Cheng Puay for help in sorting the material and helping

84THE RAFFLES BULLETIN OF ZOOLOGY 2004
with the study in various ways. This study was partially
supported by funding from the Workshop on Managing
Potential Conflicts in the South China Sea.

Ng, P. K. L. & P. J. F. Davie, 1991. The Indo-Pacific Pilumnidae
VII. Notes on the taxonomy of Heteropilumnus sasekumari(Serène, 1971) and Cryptolutea Ward, 1936 (Crustacea:
Decapoda: Brachyura). Memoirs of the Queensland Museum,30(3): 517-524.

LITERATURE CITED

Ng, P. K. L. & L. W. H. Tan, 1988. The identities of Heteropilumnus
subinteger (Lanchester, 1900) and Heteropilumnus hirsutior(Lanchester, 1900) stat. nov., with description of a new species,
Heteropilumnus holthuisi sp. nov. (Crustacea, Decapoda,
Brachyura, Pilumnidae). Crustaceana, 54(1): 13-24.

Bott, R., 1970. Die Süsswasserkrabben von Europa, Asien,
Australien und ihre Stammesgeschichte. Eine Revision der
Potamoidea und Parathelphusoidea (Crustacea, Decapoda).
Abhandlungen der Senckenbergischen Naturforschenden
Gesellschaft, 526: 1-338.

Ng, P. K. L., A. G. Ilahude, N. Sivasothi & D. C. J. Yeo, 2004.
Expedition Anambas: An overview of the scientific marine
exploration of the Anambas and Natuna Archipelago, 11-22
March 2002. In: Ng, P. K. L., D. Wowor & D. C. J. Yeo (eds.),
Scientific Results of the Anambas Expedition 2002. Raffles
Bulletin of Zoology, Supplement 11: 1-17.

Dai, A. Y. & Y. Z. Song, 1986. Intertidal crabs from Beibu Gulf
of Guangxi. Transactions of the Chinese Crustacean Society,
1: 54-62, 1 pl.
Davie, P. J. F. & A. Humpherys, 1997. New species of rhizopine
crabs (Crustacea: Brachyura) from northern Australia. Memoirsof the Queensland Museum, 42(1): 97-103.

Rahayu, D. L. & P. K. L. Ng, 2000. Some new and interesting recordsof spider crabs (Decapoda, Brachyura, Majidae) from Indonesia.
Crustaceana, 73(7): 887-890.

Haan, W. De, 1833-1850. Crustacea. In: Von Siebold, Fauna
Japonica. Sive descripto animalium, qui in itinere per Japniam,
Jussuet auspiciis superiorum, qui summum in India Batava
Imperium tenet, suscepto, annis 1823-1830 collegit, notis,
observationibus et adumbrationibus illustratit. Lugduni
Batavorum, fasc. 1-8: i-xxi+vii-xvii+ix-xvi, 244 pp, pls. 1-55,
A-Q, Amsterdam.

Sakai, T., 1935. New or rare species of Brachyura, collected by the“Misago” during the zoological survey around the Izu-Peninsula.
Science Report of the Tokyo Bunrika Daigaku, (B)2: 63-88, Pls.
6-8.
Sakai, T., 1976. Crabs of Japan and the adjacent Seas, Kodansha
Ltd., Tokyo, 725 pp, 157 pls.
Schubart, C. & P. K. L. Ng, 2000. On the identities of the raftingcrabs Cancer depressus Fabricius, 1775, Cancer squamosus
Herbst, 1790, Plagusia immaculata Lamarck, 1818, andPlagusia tuberculata Lamarck, 1818 (Crustacea: Decapoda:
Brachyura: Plagusiidae). Raffles Bulletin of Zoology, 48(2):327-336.

Herbst, J. F. W., 1782-1804. Versuch einer Naturgeschichte der
Krabben und Krebse nebst einer systematischen Beschreibung
ihrer verschiedenen Arten, Berlin und Stralsund, 3 vols.,
274+226+216 pp, 72 pls.
Lamark, M. de, 1818. Homobranches Brachyures. In: Les caractéres
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Stephenson, W., 1972. An annotated checklist and key to the IndoWest Pacific swimming crabs (Crustacea: Decapoda:
Portunidae). Bulletin of the Royal Society of New Zealand, 10:
1-64.

Man, J. G. De, 1895. Bericht uber die von Herrn Sciffscaptan Storm
zur Atejeh, an den westlichen Kuchen von Malakka, Borneo
und Celebes sowie in der Java-See gesammelten Decapoden und
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Stephenson, W. & B. Campbell, 1959. The Australian portunids(Crustacea: Portunidae). III. The genus Portunus. Australian
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Wagner, H. P., 1992. A new species of Austrolibinia from the South
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Ng, P. K. L., 1986. Preliminary descriptions of 17 new freshwater
crabs of the genera Geosesarma, Parathelphusa, Johora andStoliczia (Crustacea, Decapoda, Brachyura) from South East
Asia. Journal of the Singapore National Academy of Science,
15: 36-44.

Ward, M., 1936. Crustacea Brachyura from the coasts of Queensland.
Memoirs of the Queensland Museum, 11(1): 1-13, pls 1-3.
Yamaguchi, T. & K. Baba, 1993. Crustacean specimens collected
in Japan by Ph. F. von Siebold and H. Bürger and held by the
Nationaal Natuurhistorisch Museum in Leiden and othermuseums. In: Yamaguchi, T. (ed.), Ph. F. von Siebold andNatural History of Japan Crustacea. Carcinological Society ofJapan, Tokyo: 145-570.

Ng, P. K. L., 1987. The Indo-Pacific Pilumnidae II. A revision of
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Zoology, 4(1): 69-111, pl. 1.

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Yeo et al.: Anambas Expedition Brachyura
Appendix 1. Checklist of Crustacea Brachyura of the Anambas Expedition. For station data, see Ng et al., 2004 (in present volume).
Species treated in detail in present study under Taxonomic Notes marked by an asterisk(*).
FAMILY/SPECIESSTATION (NUMBER OF SPECIMENS, DEPOSITORY)FAMILY DORIPPIDAE MACLEAY
Subfamily Dorippinae MacLeay
Dorippe quadridens (Fabricius)

EA-TT 04 (1 female, MZB)FAMILY LEUCOSIIDAE SAMOUELLE
Subfamily Ebaliinae Stimpson
Arcania undecimspinosa De Haan

EA-TT 08 (1 male, 1 female, MZB)FAMILY MAJIDAE SAMOUELLE
Subfamily Majinae Samouelle
Cyclax spinicinctus Heller
Schizophrys aspera (H. Milne Edwards)

EA-D 11 (1 female, ZRC)
EA-D 11 (2 males, MZB; 1 male, 1 female, ZRC)

Subfamily Pisinae Dana
Austrolibinia pincerna Wagner *Hyastenus diacanthus (De Haan)
Lahaina ovata Dana
Phalangipus longipes (Linnaeus)
Tylocarcinus styx (Herbst)

EA-TT 01 (1 male, ZRC); EA-TT 06 (1 female, MZB)EA-TT 01 (1 male, MZB); EA-TT 03 (1 male, ZRC)
EA-D 11 (5 males, 2 females, MZB; 5 males, 1 female, ZRC)
EA-TT 01 (2 males, MZB); EA-TT 04 (1 female, MZB); EA-TT 08(1 male, 2 females, ZRC)
EA-D 03 (2 males, 2 females, ZRC); EA-D 11 (2 males, MZB)

Subfamily Epialtinae MacLeay
Xenocarcinus depressus Miers

EA-D 11 (4 males, 2 females, MZB; 5 males, 2 females, ZRC)

Subfamily Mithracinae MacLeay
Micippa platipes Rüppell

EA-ZJ 10 (1 female, MZB)FAMILY PORTUNIDAE RAFINESQUE
Subfamily Portuninae RafinesquePortunus (Monomia) argentatus argentatus (White)

Portunus (Monomia) rubromarginatus (Lanchester)Portunus (Portunus) pelagicus (Linnaeus)
Portunus (Xiphonectes) gracilimanus (Stimpson)Portunus (Xiphonectes) hastatoides Fabricius
Portunus (Xiphonectes) pulchricristatus (Gordon)
Portunus (Xiphonectes) tenuipes (De Haan) *Scylla olivacea (Herbst)

Subfamily Caphyrinae Paul’son
Caphyra rotundifrons (A. Milne-Edwards)
Lissocarcinus polybiodes Adams & White
Subfamily Thalamitinae Paul’son
Charybdis (Goniohellenus) truncata (Fabricius)
Thalamita admete (Herbst)
Thalamita crenata (Latreille)

Thalamita danae Stimpson

Thalamita spinicarpa Wee & Ng
Thalamitoides quadridens A. Milne-Edwards

EA-TT 01 (1 male, 1 female, MZB); EA-TT 02 (1 female, ZRC); EA-TT
06 (7 males, 9 females, MZB; 5 males, 5 females, ZRC); EA-TT 08(3 males, 2 females, ZRC)
EA-TT 01 (1 male, ZRC); EA-TT 03 (1 male, MZB); EA-TT 04 (2 males,
2 females, ZRC; 3 males, 2 females, MZB)EA-D 13 (1 juvenile MZB); EA-DW 10 (1 male, ZRC)
EA-TT 02 (1 male, MZB)EA-TT 06 (2 males, 2 females, MZB; 1 male, 1 female, ZRC); EA-TT
08 (13 males, 13 females, MZB; 5 males, 5 females, ZRC)
EA-TT 06 (1 male, MZB; 1 male, ZRC)
EA-TT 01 (3 males, 6 females, MZB); EA-TT 04 (4 males, 4 females,
ZRC); EA-TT 06 (1 female, MZB)
DW 0205 (2 males, MZB); EA-DW 02A (2 males, ZRC); EA-DW 09(1 male, MZB; 1 juvenile ZRC)EA-ZJ 09 (1 female, ZRC)
EA-TT 03 (1 female, MZB)EA-TT 06 (1 male, 1 female, MZB); EA-TT 08 (5 males, 5 females, MZB;
5 males, 2 females, ZRC)
EA-JL 01 (3 males, 1 female, MZB; 2 males, ZRC)
EA-DW 04 (1 female, MZB); EA-DW 10 (1 male, ZRC); EA-JL 01(3 males, 1 female, MZB); EA-JL 06 (1 male, 1 female, ZRC), ; EAJL 07 (4 males, 3 females, MZB); EA-ZJ 01 (1 female, MZB); EAZJ 08 (1 male, ZRC); EA-ZJ 10 (1 female, MZB); EA-ZJ 11 (2females, MZB)EA-JL 01 (1 female, MZB), 04 (1 male, ZRC), 07 (2 females, ZRC); EAZJ 01 (2 males, ZRC; 2 males, MZB), 04 (1 male, MZB), 11 (1 male,ZRC)
EA-JL 03 (1 male, MZB)EA-D 11 (1 female, MZB); EA-ZJ 12 (1 female, 1 juvenile ZRC)

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FAMILY ERIPHIIDAE MACLEAY
Subfamily Eriphinae MacLeay
Eriphia smithii MacLeay
Eriphia sebana (Shaw & Nodder)

EA-ZJ 04 (1 female, MZB)EA-ZJ 09 (1 male, 4 females, MZB); EA-ZJ 10 (1 male, 2 females, ZRC);EA-ZJ 11 (1 male, ZRC)

Subfamily Oziinae Dana
Epixanthus dentatus (White)
Ozius guttatus H. Milne Edwards
Ozius tuberculosus H. Milne Edwards

EA-ZJ 05 (1 male, 1 female, MZB)EA-JL 06 (1 female, MZB); EA-JL 07 (1 male, ZRC; 1 male, MZB)EA-ZJ 09 (1 female, ZRC)FAMILY CARPILIIDAE ORTMANN, 1893
Carpilius maculatus (Linnaeus)

EA-ZJ 04 (1 female, ZRC); EA-D 11 (1 female, MZB)FAMILY XANTHIDAE MACLEAY
Subfamily Xanthinae MacLeay
Leptodius exaratus (H. Milne Edwards)
Leptodius davaoensis Ward
Leptodius sanguineus (H. Milne Edwards)
Subfamily Actaeinae Alcock
Actaeodes hirsutissimus (Rüppell)
Actaeodes tomentosus (H. Milne Edwards)
Pseudoliomera granosimana (A. Milne-Edwards)

EA-JL 07 (2 males, 1 female, MZB); EA-ZJ 07 (1 female, ZRC); EA-ZJ
11(1 male, ZRC)
EA-JL 01 (1 male, MZB)EA-JL 07 (1 female ov., ZRC)

EA-D 04 (1 male, MZB)EA-JL 04 (1 male, 2 females, ZRC); EA-ZJ 04 (3 males, 2 females, MZB);EA-ZJ 06 (1 male, 1 female, ZRC)EA-ZJ 05 (1 female, MZB)

Subfamily Zosiminae Alcock
Lophozozymus pulchellus A. Milne-Edwards

EA-D 11 (1 female, ZRC)

Subfamily Etisinae Ortmann
Etisus laevimanus Randall

EA-JL 06 (1 female, MZB)

Subfamily Chlorodinae Dana
Cyclodius granulatus (Targioni-Tozzetti)
Pilodius nigrocrinitus Stimpson

Pilodius paumotensis Rathbun

EA-D 11 (1 female, MZB)EA-JL 04 (2 males, 1 female, ZRC); EA-JL 07 (1 male, 1 female, ZRC);EA-ZJ 01 (2 males, 1 female, MZB); EA-JL 03 (1 female, ZRC);
EA-JL04 (1 male, ZRC); EA-JL 05 (1 male, 5 females, MZB); EAJL 06 (1 male, MZB); EA-JL 07 (1 male, MZB)EA-ZJ 01 (1 male, ZRC); EA-ZJ 03 (1 female, MZB)FAMILY TRAPEZIIDAE MIERS
Tetralia nigrolineata Serène & Pham
Tetralia rubridactyla Garth
Trapezia cymodoce (Herbst)
Trapezia lutea Castro
Trapezia septata Dana

EA-D 03, EA-D 06 (1 male, 1 female, MZB)EA-D 11 (1 male, MZB)EA-D 03 (1 male, MZB)EA-D 11 (1 male, 1 female, ZRC)
EA-D 03 (1 male, 1 female, MZB); EA-D 11 (3 males, 2 females, ZRC)FAMILY PILUMNIDAE SAMOUELLE
Subfamily Pilumninae Samouelle
Actumnus setifer (De Haan)
Bathypilumnus sinensis (Gordon)
Glabropilumnus laevimanus (Dana)
Pilumnus minutus De Haan
Pilumnus scabriusculus Adams & White
Pilumnus vespertilio (Fabricius)

EA-TT 04 (1 female ov., ZRC)
EA-TT 04 (1 female ov., MZB)EA-JL 02 (2 males, 1 female, ZRC); EA-JL 07 (4 males, 4 females, MZB);EA-ZJ 05 (1 male, MZB)EA-TT 01 (1 male, 1 female, MZB); EA-TT 04 (1 male, MZB; 1 male,
ZRC); EA-ZJ 05 (1 female,MZB); EA-D 11 (1 male, MZB)EA-ZJ 05 (1 female, MZB; 1 female, ZRC)
EA-DW 04 (1 female, ZRC); EA-JL 01 (2 males, 2 females, MZB); EAJL 02 (1 female, ZRC); EA-JL 03 (1 male, 2 females, ZRC); EA-JL
06 (1 male, MZB); EA-JL 07 (1 female, ZRC); EA-ZJ 01 (1 male,
3 females, ZRC); EA-ZJ 04 (1 female, ZRC)

Subfamily Rhizopinae Stimpson
Heteropilumnus satriai, new species *EA-D 11 (1 male, MZB)

Subfamily Tanaochelinae Ng & Clark
Tanaocheles bidentata (Nobili)

EA-JL 01 (2 females, MZB); EA-JL 03 (1 female, ZRC)

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Yeo et al.: Anambas Expedition Brachyura
Subfamily Eumedoninae Dana
Zebrida adamsii WhiteFAMILY OCYPODIDAE RAFINESQUE
Subfamily Ocypodinae RafinesqueOcypode ceratophthalmus (Pallas)
Ocypode cordimanus LatreilleUca annulipes (H. Milne Edwards)
Uca forcipata (Adams & White)
Uca perplexa (H. Milne Edwards)

Uca triangularis (A. Milne-Edwards)
Uca vocans (Linnaeus)

Subfamily Macrophthalminae Dana
Macrophthalmus (Chaenostoma) boscii Audouin
Macrophthalmus (Macrophthalmus) convexus Stimpson
Macrophthalmus (Macrophthalmus) crassipes
H. Milne Edwards
Macrophthalmus (Macrophthalmus) telescopicus Owen
Subfamily Scopimerinae Alcock
Scopimera globosa (De Haan)
FAMILY GRAPSIDAE MACLEAY
Grapsus albolineatus Lamarck

Metopograpsus frontalis Miers
Metopograpsus oceanicus (Hombron & Jacquinot)

Pachygrapsus minutus A. Milne-Edwards
Pachygrapsus planifrons De Man
FAMILY VARUNIDAE H. MILNE EDWARDS
Ilyograpsus paludicola (Rathbun)
Ptychognathus barbatus (A. Milne-Edwards)
Utica borneensis De Man
Varuna yui Hwang & Takeda

FAMILY SESARMIDAE DANA
Episesarma sp. (juvenile)
Parasesarma anambas, new species *Perisesarma eumolpe (De Man)
Perisesarma indiarum (Tweedie)
Perisesarma sp.
Pseudosesarma bocourti (A. Milne-Edwards)

probably TT-06 (this specimen was photographed on site but lost
subsequently)

EA-ZJ 01 (1 male, ZRC); EA-ZJ 10 (2 males, 2 females, MZB)EA-JL 07 (1 male, MZB); EA-ZJ 03 (1 male, MZB); EA-ZJ 10 (1 male,
MZB)
EA-DW 05 (1 male, 2 females, MZB; 1 male, 1 female, ZRC)
EA-DW 04 (1 male, ZRC); EA-JL 01( 1 male, 1 female, MZB; 2 males,
1 female, ZRC),
EA-JL 01 (46 males, 40 females, ZRC); EA-JL 05 (13 males, 6 females,
MZB); EA-JL 09 (4 males, 1 female, MZB); EA-ZJ 08 (1 male, 1females, ZRC); EA-DW 04 (1 male, ZRC)
EA-JL 01 (4 males, 1 female, MZB; 4 males, 1 female, ZRC)
EA-JL 01 (2 males, 1 female, MZB); EA-JL 03 (1 male, 3 females, ZRC);
EA-JL 06 (4 males, 4 females, MZB; 1 male, 1 female, ZRC); EAZJ 01 (1 male, 5 females, ZRC)

EA-JL 01 (1female, ZRC) EA-ZJ 01 (1 male, MZB)EA-ZL 01 (2 females, MZB)EA-ZJ 08 (1 female, MZB)EA-ZJ 01 (1 female, MZB; 1 female, ZRC)

EA-DW 05 (2 males, MZB; 1 male, ZRC)

EA-JL 07 (1 male, 1 female, MZB); EA-ZJ 08 (3 males, 2 females, ZRC);EA-ZJ 09 (1 male, 6 female, ZRC); EA-ZJ 10 (4 males, 4 females,
MZB); EA-ZJ 11(1 male, ZRC)
EA-JL 02 (1 female, MZB)EA-JL 01 (12 males, 19 females, ZRC); EA-JL 02 91 female, ZRC); EAJL 03 (2 males, 2 females, ZRC); EA-JL 05 (1 male, ZRC); EA-JL
07 (2 males, 3 females, MZB); EA-ZJ 03 (4 males, 1 female, ZRC);EA-ZJ 06 (6 males, 7 females, MZB); EA-ZJ 07 (1 male, ZRC); EAZJ 08 (2 males, ZRC); EA-ZJ 10 (1 male, 1 female, ZRC); EA-ZJ 11(2 males, ZRC)
EA-JL 07 (1 female, MZB)EA-JL 07 (1 female ov., MZB)EA-DW 04 (2 females ov., ZRC)EA-ZJ 11 (1 female ov., MZB)EA-DW 04 (2 males, 3 females, ZRC)
EA-DW 02 (2 males, MZB; 1 male, 1 female, ZRC); EA-DW 03 (1 female,
MZB); EA-DW 09 (1 female, MZB)EA-JL 08 (4 females, ZRC)
EA-DW 04 (1 male, MZB); EA-JL 05 (1 female, MZB; 1 male, 2 females,ZRC)
EA-JL 01 (3 females, MZB)EA-JL 01, EA-JL 05 (3 males, 10 females, MZB)EA-JL 06, EA-JL 09 (8 females, ZRC)
EA-DW 14 (4 females, MZB)FAMILY PLAGUSIIDAE DANA
Plagusia squamosa (Herbst) *EA-ZJ 10 (1 male, MZB)FAMILY GECARCINIDAE MACLEAY
Cardisoma carnifex (Herbst) *DW 0205 (3 males, ZRC; 3 males, MZB)

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