RAFFLES BULLETIN OF ZOOLOGY 2014
Taxonomy & Systematics
RAFFLES BULLETIN OF ZOOLOGY 62: 389–395

Date of publication: 27 June 2014
http://zoobank.org/urn:lsid:zoobank.org:pub:8493E540-DB48-4CA6-8A73-9D232D5546E1

Taxonomy of the freshwater crab Coccusa cristicervix Ng & Grinang,
2004 (Decapoda: Brachyura: Gecarcinucidae), with description of a
new species of Thelphusula from Sarawak, Malaysia, Borneo
Jongkar Grinang1* & Peter K. L. Ng2
Abstract. The taxonomy of the Sarawakian gecarcinucid freshwater crab Coccusa cristicervix Ng & Grinang, 2004,
is revised with the recent discovery of males, and the species is transferred to Thelphusula Bott, 1969. The maleabdomen, first and second gonopods are described and figured for the first time. A new species, Thelphusula pueh,
is also described from swampy habitats in Kampung Sebako in Pueh, Lundu District, Sarawak.
Key words. Crustacea, Brachyura, Gecarcinucidae, taxonomy, Thelphusula, Sarawak, Malaysia

INTRODUCTION

T. pueh, from Kampung Sebako, Pueh, Lundu District, ineastern Sarawak.

The Bornean gecarcinucid genus Thelphusula Bott, 1969,
has nine recognised species: T. baramensis (De Man,
1902), T. buergeri (De Man, 1899) (= Gecarcinucus(Cylindrothelphusa) buergeri lebangensis (Balss, 1937)),
T. dicerophilus Ng & Stuebing, 1990, T. granosa Holthuis,
1979, T. hulu Tan & Ng, 1997, T. luidana (Chace, 1938),
T. sabana Tan & Ng, 1998, T. styx Ng, 1989b, and T.
tawauensis Tan & Ng, 1998 (cf. Tan & Ng, 1998; Ng etal., 2008). The distribution of the genus in Borneo appears
to be confined to the northern half of the island, with T.
buergeri found in central Kalimantan. On the other hand,the allied genus Coccusa Tan & Ng, 1998, established for C.
isophallus Tan & Ng, 1998 (type species) and C. adipis (Ng& Wowor, 1990), occurs in eastern and western Kalimantan.
The distributions of these taxa may not be real and merely aconsequence of collecting efforts, with Sarawak and Sabahbeing relatively better surveyed. Species of Thelphusula andCoccusa, however, have very cryptic habits, being nocturnaland burrowers in swampy and/or well forested areas, andhence not easily collected (Tan & Ng, 1998; pers. obs.).
More species can be expected to be discovered from Borneo.

MATERIAL AND METHODSThe terminology used follows Ng (1988), with theabbreviations G1 and G2 used for the male first andsecond gonopods, respectively. Measurements provided(in millimetres) are of the carapace width and length,
respectively. The Malay words Kampung and Gunung areused for village and mountain, respectively. The specimens
are deposited in the Sarawak Biodiversity Centre (SBC),
Kuching, Sarawak, Malaysia; and the Zoological ReferenceCollection (ZRC) of the Lee Kong Chian Natural History
Museum (ex Raffles Museum of Biodiversity Research),
National University of Singapore.
TAXONOMYFamily Gecarcinucidae Rathbun, 1904
Thelphusula Bott, 1969
Remarks. Thelphusula Bott, 1969, was established with
Potamon (Geothelphusa) buergeri De Man, 1899, as thetype species. Bott (1969) recognised five other species in thegenus as he defined it at the time: Potamon (Potamonautes)
baramensis De Man, 1902, Potamon (Geothelphusa)
hendersonianum De Man, 1899, Potamon (Geothelphusa)
kadamaianum Borradaile, 1900, Parathelphusa (Liothelphusa)
luidana Chace, 1938, and Potamon (Potamon) melanippe De
Man, 1899 (see also Bott, 1970: 58). Potamon (Thelphusa)
bidiensis Lanchester, 1900, was regarded as a juniorsubjective synonym of Thelphusula melanippe, while
Potamon kadamaianum was treated as its subspecies by Bott
(1970). Ng (1989a: 64) questioned Bott’s (1970) concept of
Thelphusula and argued that it should be separated into three
distinct morphological groups. He disputed Bott’s (1970)

Coccusa cristicervix Ng & Grinang, 2004, was described fromthe Bau area in western Sarawak on the basis of one adultfemale specimen. We have since collected more specimensof this species, and the male characters require that it betransferred to Thelphusula. We also describe a new species,

1
Institute of Biodiversity and Environmental Conservation, Universiti Malaysia
Sarawak, 94300 Kota Samarahan, Sarawak, Malaysia; Email: gjongkar@ibec.unimas.
my (*corresponding author)

Lee Kong Chian Natural History Museum, Faculty of Science, National Universityof Singapore, 6 Science Drive 2, 117543 Singapore; Email: dbsngkl@nus.edu.sg
2© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)

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synonymisation of Potamon bidiensis with T. melanippe;
recognised it as a distinct species and referred it to a
monotypic genus, Stygothelphusa Ng, 1989a. Ng (1991)
subsequently established a new genus, Arachnothelphusa, for
Potamon (Potamon) melanippe De Man, 1899, and Potamon(Geothelphusa) kadamaianum Borradaile, 1900, treating bothas distinct species. Ng (1995), in describing a new genus
from western Sarawak, showed that Potamon (Geothelphusa)
hendersonianum De Man, 1899, had been incorrectly placedin Thelphusula as it has differently structured G1 and G2, andshould be transferred to Bakousa Ng, 1995, instead. Tan &Ng (1998) revised Thelphusula, restricted the diagnosis fornine species and established a new genus, Coccusa, for two
species from Borneo in which the carapace is ovate and muchsmoother, and most significantly, the G1 terminal segment isvery long, slender and as long as the subterminal segment(versus distinctly shorter than the subterminal segment in
Thelphusula s. str.).

with somite 6 elongate, lateral margins gently concave (Fig.
1D). G1 terminal segment subconical, tip sharply tapering,
about 2 times length of subterminal segment (Fig. 3A, C–E).
G2 with distal segment very short, tip sharp (Fig. 3B).
Colour. In life, the species is red to bright red overall all
the dorsal surfaces and pereopods (adult male chelae usuallymore orangish-red), with the ventral surfaces yellowish-white(Fig. 4A, B).
Remarks. The recent series of specimens of T. cristicervix
from Sama Jaya Nature Reserve just outside Kuching city
is about 40 km from the type locality in Bau, but agree
extremely well with the holotype specimen in all aspects;
including the structure of the carapace and proportions of theambulatory legs. The good series, which includes small andlarge specimens, leaves little doubt to their conspecificity.
The morphology of the G1 of this species agrees well withwhat has been diagnosed and figured for Thelphusula Bott,
1969, by Tan & Ng (1998), with the terminal segment slender,
cylindrical and distinctly shorter than the subterminal segment(Fig. 3A, C). The carapace shape, as well as granulation of
T. cristicervix, while relatively more ovate than other known
species of Thelphusula which have more quadrate ones,
approaches the condition in more species like T. dicerophilus
and T. granosa which have relatively more inflated carapaces(cf. Holthuis, 1979: pl. 5; Ng & Stuebing, 1990: pl. 1).

Ng & Grinang (2004) recognised a third species of Coccusa
from western Sarawak. Although only one adult female
specimen was available, they noted that the general carapace
shape and features agreed better with Coccusa and therefore
referred it to this genus, even though it was the most
granulated of the three known taxa. The collection of more
specimens of the Sarawakian species, including males, showsthat Coccusa should be restricted for species with smoothercarapaces. On the basis of the gonopod structures, Coccusa
cristicervix Ng & Grinang, 2004, must be transferred to
Thelphusula instead.

In contrast to the moist karst forest where the holotype ofT. cristicervix was collected, the recent specimens were allcollected from a swampy part of a well-shaded kerangas
forest. The soil is wet and loamy, usually with a dense rootsystem permeating it. The crabs dig relatively deep burrows,
up to perhaps a metre in depth in places and come out onlyat night. The presence of the species in both limestone andkerangas forests indicates that its distribution is delimited
by soil conditions rather than forest type.

Thelphusula cristicervix (Ng & Grinang, 2004),
new combination(Figs. 1, 3A–E, 4)
Coccusa cristicervix Ng & Grinang, 2004: fig. 5; Ng, 2004: 321;
Ng & Yeo, 2007: 9; Ng et al., 2008: 69; Cumberlidge et al.,
2009: Appendix 1.
Material examined. Holotype: female (19.9 × 14.3 mm)(SBC.C.00130), Gunung Doya, limestone outcrop, Bau District,
Sarawak, coll. A. Denis et al., 2 October 2002. Others: 2 males(20.0 × 14.8 mm, 21.8 × 16.6 mm) (SBC), Sama Jaya Nature
Reserve, lowland heath swamp forest, 16 m above sea level,
Kuching Division, Sarawak, coll. J. Grinang et al., 3 July 2013; 4
males (15.1–19.4 mm × 11.3–14.8 mm), 4 females (19.1–24.8 mm
× 14.6–18.7 mm) (ZRC), Sama Jaya Nature Reserve, lowland heathswamp forest, 16 m above sea level, Kuching Division, Sarawak,
coll. P.K.L. Ng et al., 13 September 2013.

Ng & Yeo (2007: 9) noted that the species was endangered,commenting that “the Bau area [type locality] does not havelong-term protection and the species occurs in low-lying
areas, it is very vulnerable to anthropogenic and associatedeffects.” (see also Cumberlidge et al., 2009: Appendix 1).
The discovery of T. cristicervix in a nature reserve near
Kuching city does not change its status as the protectedarea is open to the public and the species is known mainly
from one small patch of swamp only several hundred square
metres in area.

Diagnosis. Carapace gently ovate; cervical groove deep,
prominent, with distinct rows of cristae running along length;
postorbital cristae low, rounded, not sharp, separated fromlow, rounded epigastric cristae low by shallow depressions;anterolateral margin entire, epibranchial teeth not discernible;
H-shaped depression relatively shallow (Fig. 1A, B).
Frontal margin deflexed; frontal median triangle not clearly
discernible, area marked by depression (Fig. 1C). Median
tooth on posterior margin of epistome subtruncate (Fig. 1C).
Ambulatory legs relatively long (Fig. 1A). Major chela stout,with gap at base of closed fingers (Fig. 1G). Male abdomen

Thelphusula pueh, new species(Figs. 2, 3F–J, 5)
Material examined. Holotype: male (16.2 × 12.1 mm)(ZRC), edge of rice field with swampy vegetation, 26 m above
sea level, Kampung Sebako, Pueh, Lundu District, Sarawak,
coll. R. Sujang, 29 November 2013. Paratypes: 6 females(12.6–19.8 mm × 9.9–15.6 mm) (ZRC), 4 females (15.8–19.4mm × 11.4–14.5 mm) (SBC), same data as holotype; 1 female(24.1 × 14.8 mm) (ZRC), in rice field, 31 m above sea level,390

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Fig. 1. Thelphusula cristicervix (Ng & Grinang, 2004). Male (19.4 × 14.8 mm) (ZRC), Sama Jaya Nature Reserve, Kuching Division,
Sarawak. A, overall dorsal view; B, dorsal view of carapace; C, frontal view of carapace; D, ventral view showing anterior thoracic
sternum and male abdomen; E, left third maxilliped; F, outer view of right chela; G, outer view of left chela.

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Fig. 2. Thelphusula pueh, new species. Holotype male (16.2 × 12.1 mm) (ZRC), Kampung Sebako, Pueh, Lundu District, Sarawak. A,
overall dorsal view; B, dorsal view of carapace; C, frontal view of carapace; D, ventral view showing anterior thoracic sternum and maleabdomen; E, left third maxilliped; F, outer view of right chela; G, outer view of left chela.

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Fig. 3. A–E, Thelphusula cristicervix (Ng & Grinang, 2004), male (19.4 × 14.8 mm) (ZRC), Sama Jaya Nature Reserve, Kuching Division,
Sarawak; F–J, Thelphusula pueh, new species, male (16.2 × 12.1 mm) (ZRC), Kampung Sebako, Pueh, Lundu District, Sarawak. A, F,
ventral view of left G1; B, G, ventral view of left G2; C, H, dorsal view of left G1; D, I, ventral view of terminal segment of left G1; E,
J, dorsal view of terminal segment of left G1. Scale bars: A–C, F–H = 1.0 mm; D, E, I, J = 0.5 mm.

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Kampung Peninjau Lama, Gunung Serambu, Bau District,
Sarawak, coll. L. Noun, 10 November 2013.

epistome subtruncate (Fig. 2C). Ambulatory legs not distinctly
elongated (Fig. 2A). Major chela with large gap between
fingers when closed (Fig. 2F). Male abdomen with somite 6
relatively short, lateral margins distinctly concave (Fig. 2D).
G1 terminal segment relatively cylindrical, straight, slightly
more than 2 times length of subterminal segment (Fig. 3F,
H–J). G2 distal segment short, curved, tip rounded (Fig. 3G).

Diagnosis. Carapace gently ovate, wider than long, dorsal
surface rugose, anterolateral margin convex, epibranchial
tooth absent, cervical groove relatively shallow, shallow
crista; postorbital cristae weak; H-shaped depression shallow(Fig. 2A, B). Frontal margin deflexed; frontal median triangle
absent (Fig. 2C). Median tooth on posterior margin ofColour. In life, the species is dark brown overall on all thedorsal surfaces, with the palm of the chela and sometimes
the legs more orange (Fig. 5A, B).
Etymology. The species is named after Pueh, the area where
it was collected. The name is used as a noun in apposition.
Remarks. Thelphusula cristicervix and T. pueh, new species,
are the only two species of Thelphusula known from western
Sarawak. They are easily differentiated from congeners inhaving a relatively more ovate carapace, a more convex
anterolateral margin and more strongly deflexed frontal
margin. Their G1s are also superficially similar in shape.
Thelphusula pueh can most easily be separated from T.
cristicervix by its live colours; with the carapace and legsof T. cristicervix red (Fig. 4A, B), even in small specimens,
whereas T. pueh is dark brown (Fig. 5A, B). Thelphusula
pueh also has proportionately shorter ambulatory legs (Fig.
2A) (versus longer in T. cristicervix, Fig. 1A); relatively
shallower cervical grooves (Fig. 2B) (versus distinctly
deeper in T. cristicervix, Fig. 1A); smooth lateral margins
of abdomen (Fig. 2D) (versus uneven in T. cristicervix, Fig.
1D); and the outer margin of the G1 subterminal segment
is almost straight with the terminal segment more strongly
bent (Fig. 3F, H) (versus outer margin of G1 subterminal
segment gradually curved with the terminal segment gently
bent in T. cristicervix, Fig. 3A, C).

Fig. 4. Colours in life, Thelphusula cristicervix (Ng & Grinang,2004). A, male (ca. 19.0 mm carapace width), in situ in forest,
specimen not collected; B, female (19.3 × 14.2 mm) (ZRC)[Photograph by: Tan HH]. Both specimens from Sama Jaya Nature
Reserve, Kuching Division, Sarawak.

Like T. cristicervix, T. pueh has a preference for damp soils
and constructs shallow burrows. Its presence in a more open
habitat (ricefields) may not be natural.
ACKNOWLEDGEMENTS
The first author thank his many friends who accompaniedand helped collect the specimens: Pui Yong Min, Tan Heok
Hui, Rowmina Sujang, Lexter Noun, and Jefry Sujang. He
is also grateful to Andrew Alek Tuen for administrative
support and Indraneil Das for informative discussions on
many aspects related to this study. Research Permit No.
NCCD.907.4.4(Jld.8)-66 was granted by Sarawak Forest
Department. The study was partially supported by Shell Chair
Grant SRC/06/2010(02); the Mohamed bin Zayed Species
Conservation Fund, Project No. 13054042; and a Raffles
Museum visiting fellowship to the first author.
Fig. 5. Colours in life, Thelphusula pueh, new species. A, paratype
female (16.3 × 12.9 mm) (SBC); B, paratype female (13.0 × 10.1mm) (SBC). Both specimens from Kampung Sebako, Pueh, Lundu
District, Sarawak.

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