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A checklist of Crustacea Decapoda collected from Conic Island Cave and adjacent areas of Hong Kong

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During October 2002, the benthic communities of Conic Island Cave and the adjacent sea area of Hong Kong were sampled by SCUBA divers. Twenty genera and 21 decapod species were identified with seven species being reported from Hong Kong for the first time.
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A checklist of Crustacea Decapoda collected from Conic
Island Cave and adjacent areas of Hong Kong
Paul F. Clark a; Peter K. L. Ng b; Charles H. J. M. Fransen c; Patsy A. McLaughlin
d; Peter C. Dworschak e; Keiji Baba f
aDepartment of Zoology, The Natural History Museum, London, UK
bDepartment of Biological Sciences, National University of Singapore, Singapore
cNationaal Natuurhistorish Museum, Leiden, The Netherlands
dShannon Point Marine Center, Western Washington University, Anacortes
eNaturhistorisches Museum Wien, Wien, Austria
fFaculty of Education, Kumamoto University, Kumamoto, Japan
Online Publication Date: 01 January 2008
To cite this Article: Clark, Paul F., Ng, Peter K. L., Fransen, Charles H. J. M.,
McLaughlin, Patsy A., Dworschak, Peter C. and Baba, Keiji (2008) 'A checklist of Crustacea Decapoda collected from
Conic Island Cave and adjacent areas of Hong Kong', Journal of Natural History, 42:9, 913 — 926
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A checklist of Crustacea Decapoda collected from Conic Island Cave
and adjacent areas of Hong Kong
Paul F. Clark
a
*, Peter K.L. Ng
b
, Charles H.J.M. Fransen
c
, Patsy A. McLaughlin
d
,
Peter C. Dworschak
e
and Keiji Baba
f
a
Department of Zoology, The Natural History Museum, London, UK;
b
Department of
Biological Sciences, National University of Singapore, Singapore;
c
Nationaal Natuurhistorish
Museum, Leiden, The Netherlands;
d
Shannon Point Marine Center, Western Washington
University, Anacortes, USA;
e
Naturhistorisches Museum Wien, Wien, Austria;
f
Faculty of
Education, Kumamoto University, Kumamoto, Japan
During October 2002, the benthic communities of Conic Island Cave and the
adjacent sea area of Hong Kong were sampled by SCUBA divers. Twenty genera
and 21 decapod species were identified with seven species being reported from
Hong Kong for the first time.
Keywords: checklist; Decapoda; Conic Island Cave; Hong Kong
Introduction
Towards the end of October 2002, a group of scientists was invited by Brian
Morton and Roger Bamber to a workshop at the Swire Marine Laboratory,
Cape D’Aguilar, Hong Kong to investigate the fauna of Conic Island Cave,
Hong Kong. SCUBA divers entered the cave to sample the benthic communities and
the epifauna on the side and roof of the cave. Sea areas adjacent to the cave were also
sampled. Sixty-two decapods were collected during this survey. These were
identified, assigned to twenty genera and twenty-one species. Seven species, five
brachyurans and two anomurans, are reported from Hong Kong for the first time.
The purpose of this paper is to provide a checklist of the decapods collected during
the survey.
Specimens examined are deposited in the Natural History Museum (NHM),
London; Zoological Reference Collection (ZRC) of the Raffles Museum, National
University of Singapore; Nationaal Naturhistorisch Museum (ex Rijksmuseum van
Natuurlijke Historie, RMNH), Leiden; Naturhistorisches Museum in Wien
(NHMW), Austria; Zoological Museum of the University of Copenhagen
(ZMUC), Denmark. Abbreviations used: CL5carapace length, coll.5collected,
juv.5juvenile, ovig.5ovigerous, PL5the length of the major cheliped, TL5total
length of specimen from tip of rostrum to the end of telson.
*Corresponding author. Email: p.clark@nhm.ac.uk
Journal of Natural History
Vol. 42, Nos. 9–12, March 2008, 913–926
ISSN 0022-2933 print/ISSN 1464-5262 online
#2008 Taylor & Francis
DOI: 10.1080/00222930701850570
http://www.informaworld.com
Downloaded By: [Natural History Museum] At: 08:32 13 June 2008
Suborder DENDROBRANCHIATA
Family PENAEIDAE
Penaeidae sp.
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 juv. (RMNH D 50576),
coll. 22 October 2002.
Remarks
The specimen is small and therefore it is not possible to assign it to a genus with any
accuracy.
Infraorder CARIDEA
Family ALPHEIDAE
Alpheus hippothoe De Man, 1888
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 ovig. R(CL. 11.4 mm
missing major chela), coll. 22 October 2002, 12.4m; 1 ovig. RCL. 6.9 mm (NHM
2004.2571), coll. 25 October 2002, 11.1m.
Hoi Ha Wan, 22u28947.430N 114u20908.80E, Hong Kong; 1 specimen, CL.
2.8 mm (NHM 2004.2572), coll. 26 October 2002, 16 m.
Remarks
This species is known from the Red Sea and East Africa to Japan, Philippines,
Indonesia and Australia, usually found in shallow water in the interstices of dead
coral.
Alpheus rapacida De Man, 1908
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 specimen (CL. 3.1 mm),
(RMNH D 50577), coll. 25 October 2002.
Hoi Ha Wan, 22u28947.43.0N 114u20908.80E, Hong Kong; 1 specimen, coll. 26
October 2002, 16m.
Remarks
This alpheid is part of the A. brevirostris species group that has no transverse groove
on the upper border of the palm of the large chela. Its major chela is about four times
longer than broad, the carapace is smooth, and the third, fourth and fifth pereiopods
have long spatulate dactyli. In the key to the Indo-West Pacific alpheid shrimps of
the brevirostris species group provided by Bruce (1994, 23), the specimen keys out
914 P.F. Clark et al.
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with A. rapacida De Man, 1908. Alpheus rapacida is known throughout the tropical
Indo-West Pacific, living in self-made burrows on sandy and muddy bottoms in
depths of 5 to 56 m.
The present specimen differs from the description of A. rapacida by De Man
(1908, 1911) in having the first and second carpal segments of the second pereiopods
of equal length while in A. rapacida the second carpal segment is 1.2 times longer
than the first.
Synalpheus sp.
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 2 juv. specimens, coll. 21
October 2002.
Synalpheus hastilicrassus Coutie`re, 1905
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 5 specimens (NHM
2004.2576), coll. 25 October 2002; 1 specimen (RMNH D 50578), coll. 25 October
2002; 2 specimens (NHM 2004.2578), coll. 25 October 2002.
Remarks
Widely distributed in the Indo-West Pacific in depths between a few and 100 metres.
Occurs in interstices of living coral heads and living in sponges.
Synalpheus neomeris (De Man, 1897)
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 6 specimens (26RMNH
D 50579; 46NHM 2004.2573), coll. 22 October 2002.
Remarks
This species is known throughout the Indo-West Pacific in depths of a few to over
100 metres. It is usually found living in association with alcyonarians, but is also
found with bryozoans and sponges.
Synalpheus streptodactylus Coutie`re, 1905
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 3 specimens (16RMNH
D 50580; 26NHM 2004.2574), coll. 21 October 2002; 1 RCL 3.4 mm (abdomen
partly broken off), (NHM 2004.2577), coll. 25 October 2002.
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Remarks
Widely distributed in the Indo-West Pacific, occurring in shallow water. Associated
with sponges.
Family PALAEMONIDAE
Palaemonella rotumana (Borradaile, 1898)
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 2 specimens (RMNH D
50581), coll. 21 October 2002; 1 specimen (NHM 2004.2569), coll. 22 October 2002;
1 specimen (NHM 2004.2570), coll. 25 October 2002.
Remarks
Palaemonella rotumana is a browser found inside sponges, and its presence here in
and around the cave epifauna is not unexpected. This species is widely distributed in
the Indo-West Pacific.
Family HIPPOLYTIDAE
Thor paschalis (Heller, 1862)
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 juv. (NHM 2004.2575),
coll. 25 October 2002.
Remarks
This genus is widely distributed in shallow waters of the Indo-West Pacific. The
present specimen is small but can be identified to species.
Infraorder THALASSINIDEA
Family CALLIANASSIDAE
Subfamily CALLIANASSINAE
(sensu Manning & Felder, 1991)
Callianassa sp.
Material examined
Long Kei Wan, 22u219540N 114u239220E, Hong Kong; 1 ? (NHM 2003.1058), 1 R,
13.7 mm (NHMW 19899), coll. 25 October 2002.
Remarks
Both specimens are small, but the larger specimen (TL55.3 mm, CL51.5 mm) has a
small rostral projection, uniramous pleopod1 and biramous pleopod2, the female
gonopore being present. The two detached chelipeds (PL50.4 mm right,
916 P.F. Clark et al.
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PL50.27 mm left) probably belong to this specimen. The smaller specimen
(TL54.1 mm, CL51.1 mm) has almost no rostrum projection and lacks pleopod1+2
and a female gonopore (so the sex is not known).
All chelipeds have a median meral spine, with small spines on the cutting edges of
the fixed finger and the ischium is unarmed. The eyestalks are relatively large, with
the cornea distinct and almost terminal in position (juvenile). The rostral projection
is relatively short in the larger specimen and not visible in the smaller specimen. The
third maxilliped is operculiform, with the merus slightly projecting beyond the
articulation, the crista dentata with only a few spines (juvenile condition). The
antenna peduncle is slightly longer than the antennule, with the antennular flagellum
shorter than the peduncle. The specimens are juveniles (many characters are not yet
fully developed: pleopods1+2, eyestalks, rostral projection, spination of pereiopod1
merus etc.) and too small to be identified.
Even assignment to one of the genera recognised by Manning and Felder (1991)
and Poore (1994) for these specimens is difficult for the reason given above, and the
fact that some genera are also weakly defined. As things remain, the specimens key
out to Biffarius in Poore (1994) but do not fit exactly, as they will probably develop
longer rostral projections when larger. The alternative would be Pseudobiffarius,a
genus erected by Heard and Manning (2000), but specimens do not fit exactly with
regards to the form of the antennae and pleopods1+2. Sakai (1999b) synonymised
Biffarius (and all genera attributed to the subfamily Callianassinae) with Callianassa.
However, Callianassa [sensu lato] is considered to be a polyphyletic clade, with
species included as listed in Tudge et al. (2000), but not in the sense of Sakai (1999b)
– see Ngoc-Ho (2003) and Dworschak (2003). In any case the genera within the
Callianassidae still require revision.
Infraorder ANOMURA
Family PAGURIDAE
Pagurus hedleyi (Grant & McCulloch, 1906)
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 specimen (NHM
2003.971), coll. 22 October 2002.
Remarks
Pagurus hedleyi and P. kulkarnii Sankolli, 1962 look very much alike and the small
size of the present specimen and absence of colour make its precise identification
difficult. Although both occur in Hong Kong, the characters of the present specimen
more likely identify it as P. hedleyi.
Pagurus imaii (Yokoya, 1939)
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 specimen (NHM
2003.970), coll. 22 October 2002.
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Remarks
The specimen here is a very small female. To date this species is known only from
Japan and this is therefore the first record from Hong Kong. Nevertheless, the
combination of characters of the present specimen show that it is P. imaii as
presently understood. Komai (1999) transferred Pagurus imaii to Parapagurodes, but
Mclaughlin and Asakura (2004) referred it back to Pagurus.
Anapagrides sp.
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 specimen (NHM
2002.968), coll. 22 October 2002.
Remarks
The present specimen is a male with a regenerating right cheliped. While reasonably
confident of the generic placement, this can only be confirmed with a female; and
more specimens will be needed to ascertain the identity of the species.
Family PORCELLANIDAE
Petrolisthes boscii (Audouin, 1826)
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 specimen (NHM
2003.974), coll. 22 October 2002.
Remarks
Reported from Hong Kong by Haig (1992). Widely dispersed from Red Sea, Persian
Gulf, Thailand, Hong Kong, Japan, Australia and Fiji Islands.
Pisidia gordoni Johnson, 1970
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 specimen (NHM
2003.977), 21 October 2002; 1 specimen (NHM 2003.973), coll. 22 October 2002.
Hoi Ha Wan, 22u28947.430N 114u20908.80E, Hong Kong; 1 specimen (NHM
2003.975), coll. 26 October 2002, 12m.
Remarks
The specimen collected on 22 October has no carapace, antennae or antennules, and
its placement here is tentative. Reported from Hong Kong by Haig (1992). Widely
reported from Red Sea, East Africa, Arabian Gulf, Hong Kong, Thailand,
Singapore and Australia.
918 P.F. Clark et al.
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Pisidia serratifrons (Stimpson, 1858)
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 specimen (NHM
2003.976), coll. 21 October 2002.
Remarks
Reported from Hong Kong by Haig (1992). Reported from Korea, China and
Japan.
Pisidia cf. dispar (Stimpson, 1858)
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 specimen (NMH
2003.972), coll. 22 October 2002.
Remarks
There is only one very small ovigerous female that has only one cheliped and one
ambulatory leg left. As such, its identity here is tentative. Reported from Hong Kong
by Haig (1992). Distribution from Hong Kong to Japan, southwards Australia and
east to Fiji.
Enosteoides ornatus (Stimpson, 1858)
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 specimen (NHM
2003.969), coll. 25 October 2002.
Remarks
Only one tiny specimen, probably a juvenile, is available for study. Reported from
Hong Kong by Haig (1992). Distribution from Korea to Japan, Singapore to
Australia.
Family GALATHEIDAE
Galathea ohshimai Miyake & Baba, 1967
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 R(NHM 2004.305), coll.
21 October 2002.
Remarks
This female is the first record of the species from Hong Kong. It was previously
recorded from the Palau Islands (type locality), Moluccas and New Caledonia.
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Galathea orientalis Stimpson, 1858
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 R(NHM 204.304), coll.
21 October 2002; 1 (NHM 2004.303), coll. 25 October 2002.
Remarks
This species was first described from Hong Kong, and is now known from Japan
including the Bonin islands, Korea, East China Sea including Tawain, South China
Sea, and Western Australia. The material reported by Tirmizi (1966) from the Red
Sea and South Arabian Sea may in all probability be identical with G. anepipoda
Baba, 1990.
Infraorder BRACHYURA
Family MAJIDAE
Hyastenus hilgendorfi De Man, 1887
Material examined
Hoi Ha Wan, 22u28947.430N 114u20908.80E, Hong Kong; 1 (7.565.3 mm) (NHM
2003.696), inside epifauna, coll. 26 October 2002, 12 m.
Remarks
This is a common species with a wide distribution from the western Indian Ocean to
Hawaii (Griffin and Tranter 1986; 147), although it has not been previously recorded
from Hong Kong. The gonopod1 structure of the male specimen is diagnostic (cf.
Griffin and Tranter 1986; Fig. 42h, I).
Family ERIPHIIDAE
Sphaerozius nitidus Stimpson, 1858
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 R(11.668.5 mm)
(NHM 2003.698), coll. 22 October 2002.
Remarks
This is a well known species associated with fouling organisms, preferring areas with
relatively fast flowing waters. Hong Kong is its type locality (see also Stimpson
1907). Only one species is currently recognized in the current literature, S. nitidus
Stimpson, 1858, in Sphaerozius Stimpson, 1858, but indications are that there are at
least two taxa. Examination of the syntypes of one of the purported synonyms,
Cancer scaber Fabricius, 1798, in ZMUC (1 , 15.5612.3 mm, Cru 112-4; 1 ,
18.0614.0 mm, Cru 112-1; 1 R, 18.4614.2 mm, Cru 112-2; 1 R, 18.4614.2 mm,
920 P.F. Clark et al.
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112-3; 1 R, 17.7613.5 mm, Cru 112-5; ‘‘India Orientali’’, Dom. Daldorff) suggests it
is a distinct species, distinguished most easily by its far more granulose carapace. For
this present study a good series of specimens was examined and these are referable to
S. nitidus, from Singapore, Malaysia and southern China. Sakai (1999a, 31)
recognised Sphaerozius panope (Herbst, 1801) as a distinct species in his reappraisal
of Herbst’s taxa, but from his figure (Sakai, 1999a: pl. 16C), it is clear that it is the
same as Sphaerozius scabrus (Fabricius, 1798). Pilumnopeus granulosus Miers, 1880,
is also probably the same taxon.
With regard to the other junior synonyms of S. nitidus,viz.Actumnus nudus A.
Milne-Edwards, 1867, Menippe ortmanni De Man, 1899, Menippe convexa Rathbun,
1894, and Sphaerozius oeschi Ward, 1941, all appear to be identical with what is here
recognized as S. nitidus. In any case, a revision is clearly necessary
Family XANTHIDAE
Epiactaea nodulosa (White, 1848)
Material examined
Hoi Ha Wan, 22u28947.430N 114u20908.80E, Hong Kong; 1 (7.765.3 mm) (NHM
2003.699), coll. 26 October 2002, 12 m; 1 (12.168.6 mm) (NHM 2003.697).
Hoi Ha Wan, 22u23947.200N 114u20908.70E, Hong Kong; 1 (11.568.2 mm)
(ZRC 2003.0345), coll. 26 October 2002, 16 m.
Remarks
The genus Epiactaea Sere`ne, 1984, was established for three species, Actaea bullifera
Alcock, 1898, Actaea margaritifera Odhner, 1925, and Actaea nodulosa White, 1848 (type
species). The most often reported is E. nodulosa (Actaea pisigera Nobili, 1905, is a
subjective synonym) which has been recorded from various parts of the Indian Ocean
and Southeast Asia. Its presence in Hong Kong and China is, however, new.
Genus Pilumnidae
Actumnus elegans De Man, 1888
Material examined
Conic Island Cave, 22u219540N 114u239220E, Hong Kong; 1 R(6.665.0 mm) (ZRC
2003.0344), coll. 25 October 2002; 1 young R(4.563.5 mm) (NHM 2003.700), coll.
21 October 2002.
Remarks
This is a species typically associated with epifauna and sponges, often in areas with
faster flowing waters (see Goh et al. 1990). The rounded carapace with the low and
serrated anterolateral lobes, as well as the simple gonopod1 with a straight distal
part, and truncate tip are diagnostic characters (Takeda and Miyake 1969). Its
presence in Hong Kong is recorded for the first time. This species may need assigning
to its own genus.
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Nanopilumnus rouxi (Balss, 1935)
Material examined
Hoi Ha Wan, 22u28947.430N 114u20908.80E, Hong Kong; 1 (3.762.6 mm) (NHM
2003.701), coll. 26 October 2002, 12 m.
Remarks
Takeda (1974) established Nanopilumnus for six species previously placed in other
genera, with Medaeus rouxi Balss, 1935, as the type species. Nanopilumnus rouxi has
been reported from India, Singapore, Japan and now Hong Kong.
Latopilumnus conicus Ng & Clark, 2008
Material examined
Conic Island Cave, 22u219540N114u239220E, Hong Kong, 2 „„ (3.962.8 mm,
3.362.3 mm), 2 RR (4.863.3 mm, 5.063.4 mm) (ZRC 2003.0343), coll. 25 October
2002; 1 ovig. R(3.262.2 mm) (NHM 2003.689), coll. 21 October 2002; 1
(3.862.8 mm), 2 RR (4.062.8 mm, 3.362.3 mm) (NHM 2003.692–694), coll. 21
October 2002; 1 (3.662.6 mm) (NHM 2003.695), coll. 22 October 2002; 1
(4.963.3 mm), 1 ovig. R(3.662.7 mm) (NHM 2003.690–691), coll. 21 October 2002;
1juv. (NHM 2003.978), coll. 25 October 2002.
Remarks
This is a new species for Hong Kong. The detailed taxonomy of this species is treated
elsewhere in this volume (see Ng and Clark 2008).
Discusssion
The present records from Conic Island are interesting in that several of them (Pisidia
serratifrons (Stimpson, 1858), Enosteoides ornatus (Stimpson, 1858), Galathea
orientalis Stimpson, 1858, Sphaerozius nitidus Stimpson, 1858) are species which
were originally described from or near Hong Kong and important for future
taxonomic work, especially since most of Stimpson’s (1858a,b) types are no longer
extant. Also of interest is the fact that the anomuran and brachyuran crab fauna of
Hong Kong is still not that well studied, with seven of the specimens collected,
Hyastenus hilgendorfi De Man, 1887, Epiactaea nodulosa (White, 1848), Actumnus
elegans De Man, 1888, Nanopilumnus rouxi (Balss, 1935), Latopilumnus malardi (De
Man, 1913), Pagurus imaii (Yokoya, 1939) and Galathea ohshimai Miyake & Baba,
1967, being new records.
None of the species treated here can be considered to be a true cavernicolous
dweller, and all have been previously reported from more open habitats. A good
number of them, however, are clearly closely associated with fouling communities as
well as being interstitial occupants of sponges and rocks; this suggests that this habit
would deserve more attention in future surveys of Hong Kong. Certainly, much of
the earlier published work on Hong Kong decapod crustaceans from subtidal waters
922 P.F. Clark et al.
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has been based on the collections of trawlers and from diving in more exposed
habitats.
Acknowledgements
P.F.C. thanks Brian Morton and Roger Bamber for arranging funds, fieldwork and
publication of the results.
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926 P.F. Clark et al.
... The species occurs on both acroporid and pocilloporid scleractinian corals as well as free-living in a wide variety of habitats from coral reef to rocky, fine sand and muddy bottoms, among seagrasses and sponges, and cave epifauna; extending from shore pools to 120 m depth (Bruce 1970;Marin et al. 2004;Li and Bruce 2006;Clark et al. 2008). ...
... While this species is known as a symbiont of corals (Bruce, 1970), it is also found on various substrates such as with Halimeda, seagrasses, hard, fine sand bottoms, and on sponges and coral heads at the depths of 0-120 m (Li and Bruce, 2006). Moreover, it is found as an endobiont species inside sponges (Clark et al., 2008). Bruce (1970) found the specimens of this species at a depth of 128 m in the southern China Sea. ...
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... The present form also shares some resemblance with H. hilgendorfi De Man (1887-1888, one recorded from Hong Kong by Clark et al. (2008), but differs from this named species by presence of an acute spine on anterior slope of gastric elevation, pronounced intestine spine, and lack of submarginal tubercles above the base of cheliped (Figs. 40a, c; see De Man (1887-1888, Griffin (1968), Griffin and Tranter (1986) Stimpson (1907), Sakai (1938) (as A. spinifrons), Griffin (1970), Dai and Chen (1992). ...
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... More telling also is that P. convexus is not known from Southeast Asia (see above). On the basis of the many above-mentioned studies, 22 species of Pilumninae are now known from Hong Kong, most of which are intertidal and shallow-water forms (KJHW et al., unpublished data.; see also Stimpson, 1858Stimpson, , 1907Shen, 1940;Horikoshi & Takeda, 1982;Morton & Morton, 1983;Clark et al., 2008). Pilumnopeus convexus is not known to be associated with any commercial species (live seafood or ornamental), but is only known from fouling communities. ...
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The genus Galathea is one of the most speciose and unwieldy groups in the family Galatheidae. The examination of more than 9000 specimens of 144 species collected in the Indian and Pacific Oceans using morphological and molecular characters, has revealed the existence of 92 new species. The specimens examined during this study were obtained by various French expeditions supplemented by other collections from various sources, and including the type specimens of some previously described species. Most of the new species are distinguished by subtle but constant morphological differences, which are in agreement with molecular divergences of the mitochondrial markers COI and/or 16S rRNA. Here, we describe and illustrate the new species and redescribe some previously described species for which earlier accounts are not sufficiently detailed for modern standards. Furthermore we include a dichotomous identification key to all species in the genus from the Indian and Pacific Oceans.
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