Grinang et al.: A new species of Arachnothelphusa from northern Sarawak
Taxonomy & Systematics
RAFFLES BULLETIN OF ZOOLOGY 63: 454–460

Date of publication: 16 October 2015
http://zoobank.org/urn:lsid:zoobank.org:pub:4CC4DC84-8F6E-4524-9D3F-BAA3C2DAF588

A new species of tree-hole dwelling freshwater crab of the genus
Arachnothelphusa Ng, 1991 (Crustacea: Decapoda: Brachyura:
Gecarcinucidae) from northern Sarawak, Malaysia, Borneo
Jongkar Grinang1*, Pui Yong Min1 & Peter K. L. Ng2
Abstract. A new species of tree-hole gecarcinucid freshwater crab, Arachnothelphusa merarapensis, is describedfrom a primary dipterocarp forest near Merarap Hot Spring Resort in Lawas, northern Sarawak, Malaysia. This
brings the number of species in the genus Arachnothelphusa found in Borneo to five, and it is likely that more
species will be discovered from this island. The new species has preference for living in water-filled tree-holes,
which is the first record of a tree-hole crab for Southeast Asia. The habitat characteristics and distribution of thisnew species are also discussed.
Key words. Brachyura, Gecarcinucidae, Arachnothelphusa, tree-hole, Sarawak, Borneo

INTRODUCTION

MATERIAL AND METHODS

Ng (1989, 1991) reviewed the gecarcinucid genus Thelphusula
Bott, 1969, and transferred a number of long-legged
semi-terrestrial species from Borneo to two new genera,
Stygothelphusa Ng, 1989, and Arachnothelphusa Ng, 1991
(see also Tan & Ng, 1998; Ng, 2013; Ng & Grinang, 2014;
Grinang & Ng, 2014). Members of Stygothelphusa arecavernicolous in habit, have reduced body pigmentation,
proportionately longer ambulatory legs, a relatively squarish
carapace outline, and distinctly structured male first andsecond gonopods. This genus currently includes four species
viz. S. bidiense (Lanchester, 1900), S. nobilii (Colosi, 1920),
S. cranbrooki Ng, 2013, and S. antu Ng & Grinang, 2014
(Ng, 1989, 2013; Ng & Grinang, 2014). In comparison, most
species of Arachnothelphusa are free living, have relatively
shorter ambulatory legs, a more transversely ovate carapace
outline, and differently proportioned male first and second
gonopods. This genus currently contains four species, viz.
Arachnothelphusa melanippe (De Man, 1899) [central
Kalimantan], A. kadamaiana (Borradaile, 1900) [northern
Sabah], A. rhadamanthysi Ng & Goh, 1987 [eastern Sabah],and A. terrapes Ng, 1991 [eastern Sabah]. Interestingly,
Stygothelphusa has only been recorded from limestone
caves in western Sarawak, whereas Arachnothelphusa ismore widely distributed in Borneo. Here we describe a fifth
species of Arachnothelphusa, A. merarapensis, new species,
the first member of the genus to be recorded from Sarawak.

The terminology used essentially follows Ng (1988), withthe abbreviations G1 and G2 used for the male first andsecond gonopods, respectively. Measurements provided(in millimetres) are of the carapace width and length,
respectively. Specimens examined are deposited in the
Sarawak Biodiversity Centre (SBC), Kuching, Sarawak,
Malaysia; the Senckenberg Museum (SMF), Frankfurt am
Main, Germany; and the Zoological Reference Collection(ZRC) of the Lee Kong Chian National History Museum(ex Raffles Museum of Biodiversity Research), National
University of Singapore.
TAXONOMYFamily Gecarcinucidae Rathbun, 1904
Arachnothelphusa merarapensis, new species(Figs. 1–3)
Material examined. Holotype: male (22.5 × 16.8 mm)(ZRC), water-filled tree-hole, ca. 100 cm above ground,steep dipterocarp forest, Merarap Hot Spring Resort,
Lawas, northern Sarawak, Malaysia, Borneo, 4o22’25.4”N
115o26’10.1”E, 485 m asl, coll. J. Grinang & Y.M. Pui,
31 October 2014. Paratype: 1 female (19.9 × 15.2 mm)(SBC.C.00376), water-filled hole of tree buttress, ca. 90
cm above ground, steep dipterocarp forest, Merarap Hot
Spring Resort, Lawas, northern Sarawak, Malaysia, Borneo,
4o22’16.5”N 115o26’12.4”E, 494 m asl, coll. J. Grinang &Y.M. Pui, 1 November 2014; 1 female (22.3 × 16.1 mm)(SBC.C.00377), water-filled hole of tree buttress, ca. 30
cm above ground, same data as paratype, coll. Y.M. Pui,
27 February 2013.

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

Lee Kong Chian Natural History, Faculty of Science, National University of Singapore,
6 Science Drive 2, 117543 Singapore.
2© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)

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Fig. 1. Arachnothelphusa merarapensis, new species, holotype male (22.5 × 16.8 mm) (ZRC), Merarap Hot Spring, Lawas, Sarawak. A,
overall dorsal view of habitus; B, dorsal view of carapace; C, frontal view of carapace; D, ventral view showing anterior thoracic sternum
and male abdomen; E, outer view of right fourth ambulatory leg; F, left third maxilliped showing exopod; G, outer view of right chela;H, outer view of left chela.

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Grinang et al.: A new species of Arachnothelphusa from northern Sarawak

Fig. 2. Arachnothelphusa merarapensis, new species, holotype male (22.5 × 16.8 mm) (ZRC), Merarap Hot Spring, Lawas, Sarawak. A,
dorsal view of left G1; B, dorsal view of distal part of left G1; C, dorsal view of left G2; D, ventral view of left G1; E, ventral view of
distal part of left G1. Scale bars = 0.5 mm.

Comparative material. Arachnothelphusa kadamaiana(Borradaile, 1900): Holotype female (18.9 × 13.5 mm) (SMF4281), Kadamian River, Sabah, Malaysia, Borneo; 1 male
(20.1 × 14.9 mm) (SMF 4282), same data as holotype; 1
female (23.2 × 17.1 mm) (ZRC 2009.0094), Poring, Basin
1A, Sabah, Malaysia, Borneo, coll. R.F. Inger et al., 12
August 1992.

concave (Fig. 1D). G1 slender, sinuous, gently curving
outwards; terminal segment cylindrical, tapering, about halfof length of subterminal segment (Fig. 2A, B, D, E). G2
with short distal segment, about a quarter length of basal
segment (Fig. 2C).
Size. All species of Arachnothelphusa are relatively small,with the largest recorded male a freshly moulted specimenof A. terrapes measuring 25.7 mm by 18.6 mm (Ng, 1991).
The holotype male of A. merarapensis, new species, is 22.5
mm by 16.8 mm.

Diagnosis. Carapace surface convex, rugose, finely granular;
regions distinct, branchial regions slightly inflated; epigastricand postorbital cristae distinct; cervical and H-shaped gastricgrooves deep, not confluent; anterolateral margins convex,
serrated; antero- and posterolateral regions prominently
rugose, covered with numerous coarse granules; epibranchial
tooth acutely triangular, distinctly separated from external
orbital angle by wide, deep cleft; external orbital angle acutely
triangular, outer margin slightly concave, distinctly serrated(Figs. 1A–C, 3). Ischium of third maxilliped rectangular,
oblique median sulcus deep (Fig. 1F). Outer surfaces ofchelipeds distinctly rugose; carpus rugose, with granules,
inner angle with broadly triangular tooth; cutting edge ofpollex with larger median tooth (Fig. 1A, G, H). Ambulatory
legs long; merus unarmed, dorsal margin gently serrated;outer surface rugose; fourth ambulatory merus subequal tolength of carapace (Fig. 1A, E). Male abdomen T-shaped,
lateral margins of somites 5 and 6 distinctly concave; somite
6 subequal to length of telson; lateral margins of telson

Variation. The two female paratypes agree well with the
holotype male, differing only in minor non-sexual characters:
the cervical and H-grooves are proportionately deeper in thelarger female, and the ambulatory legs slightly longer. The
antero- and posterolateral regions also have relatively morecoarse granules in the larger female specimen.
Etymology. The species is named after Merarap, the type
locality where it was collected.
Colour. In life the carapace, chelipeds, and walking legs
are bright purple; the ventral surfaces of the ambulatory
legs and chelipeds are lighter coloured, and the distal partsof the fingers are cream-white (Fig. 3A, B).

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Remarks. The high density of granules and roughness ofthe dorsal carapace surface of A. merarapensis (Fig. 1A,B) easily distinguishes it from its other congeners whichhave a relatively smoother dorsal carapace surface covered
with small granules and scattered striae (cf. Fig. 5; De Man,
1899: pl. 9 fig. 11; Ng & Goh, 1987: pl. 3A; Ng, 1991: figs.
1, 3–5 [incorrectly labelled as fig. 5 with next two figure
captions not printed]). The acutely triangular external orbital
tooth separated by a wide and deep cleft is diagnostic for A.
merarapensis (Fig. 1A, B). In A. melanippe the epibranchial
tooth is distinct but relatively small and separated from thebroadly triangular external orbital tooth by a small cleft(cf. De Man, 1899: pl. 9 fig. 11; Ng, 1991: fig. 1); in A.
kadamaiana the epibranchial tooth is low and the external
orbital tooth is broad, with these two teeth appearing almost
confluent with hardly any indication of a cleft (Figs. 5A, 6A);
in A. rhadamanthysi the external orbital and epibranchial
teeth are both relatively broad due to the wide carapace (Ng& Goh, 1987: pl. 3A, B); and in A. terrapes there is a deep
and broad U-shaped cleft separating the epibranchial toothfrom the obtuse and truncate external orbital angle (Ng,1991: fig. 3 [incorrectly printed as fig. 5]). The terminal
segment of the G1 is long in A. merarapensis, about half
the length of the subterminal segment (Fig. 2A, B, D, E); it
is about one-third the length of the subterminal segment in
A. melanippe (cf. Ng, 1991: fig. 2C–F), and only a quarterof the length of the subterminal segment in A. kadamaiana
and A. terrapes (Fig. 6C–F; Ng, 1991: fig. 6D–G). The G1
terminal segment of A. terrapes is also distinct from allcongeners in that it is gently curved upwards (Ng, 1991: fig.
6D–G). The G1 of A. rhadamanthysi is not known because
no males of this species have been collected as yet.

Fig. 3. The colour in life of Arachnothelphusa merarapensis, newspecies, holotype male (22.5 × 16.8 mm) (ZRC), Merarap Hot
Spring, Lawas, Sarawak. A, dorsal view; B, frontal view.

11), until the present report there has been no evidence thatthese freshwater crabs are arboreal and live in tree holes.
Arachnothelphusa merarapensis, new species, is therefore
the first species of this genus known to be a true phytotelm
crab. All known Arachnothelphusa species seemed to display
a discrete distribution, but future molecular studies are needed
to enhance our knowledge on their phylogeographic patternin this region.

The live colours of A. melanippe and A. kadamaiana are notknown. Arachnothelphusa terrapes has brown to reddishbrown carapace, chelipeds, and walking legs (Ng, 1991: figs.
4, 5 [erroneously not labelled in original paper]) while A.
rhadamanthysi has a straw yellow coloured carapace andpereopods (Ng & Goh, 1987: 326). The live colour of thecarapace and pereopods of A. merarapensis is bright purple(Fig. 3A, B).

Habitat. The habitat of A. merarapensis is a steep primary
dipterocarp forest with a closed canopy, near the thermal
springs area of the Merarap Hot Spring Resort. The holotype
male was caught from a water-filled tree-hole, ca. 22 cm
deep, ca. 15 cm in diameter, and ca. 100 cm above ground(Fig. 4). The bottom layer of the tree-hole consists offinely decomposed organic material (plant and arthropod
debris), with the other two-thirds filled with water which
increases after heavy rain. The two other specimens werecaught from the water-filled crevices of tree buttresses.
One buttress-crevice had a diameter of about 26 cm, was
60 cm deep, and 90 cm above the ground, and contained
tightly compacted roots and leaf litter, with a hole dug into
this that was apparently constructed by the crab. The otherbuttress-crevice runs along the side of the buttress and isabout 30 cm above ground, shallow (15 cm), horizontal,
and 30 cm deep. For two consecutive nights, the first two
authors examined more than 15 tree-holes and crevices ofdifferent form, size, and conditions (either water-filled, dry,or debris-filled tree-holes), but we were only able to locate
three specimens. This species is nocturnal and highly sensitive
to light. We did not find any burrows on the soil aroundthe tree-holes, suggesting the crab is a tree-hole specialist.

The taxonomy of genus Arachnothelphusa has been discussed
at length by Ng (1991). One poorly known Sarawakian
species provisionally included in the genus by Ng (1991),
Parathelphusa (Liothelphusa) nobilii Colosi, 1920, was latershown to be a species of Stygothelphusa Ng, 1989 (see Ng &Álvarez, 2000; Ng, 2013). The new species, A. merarapensis,
described here is clearly member of Arachnothelphusa
because it has long ambulatory legs, a transversely ovate
carapace, convex anterolateral margins, rugose antero- andposterolateral carapace regions, a slender G1 whose terminal
segment is elongated and at least a quarter the length of thesubterminal segment, and an elongate G2 with a short distalsegment (Ng, 1991: 2).
All species of Arachnothelphusa are semi-terrestrial in habit
and have been recorded from limestone caves and highelevation habitats (Ng, 1991). Although a specimen of one
species, A. terrapes, was found on a tree stump (Ng, 1991:
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Grinang et al.: A new species of Arachnothelphusa from northern Sarawak

Fig. 4. Illustration of Arachnothelphusa merarapensis, new species,
holotype male (22.5 × 16.8 mm) (ZRC), emerging at night fromits filled-water tree-hole in primary dipterocarp forest, Merarap
Hot Spring, Lawas, Sarawak.

Fig. 5. Arachnothelphusa kadamaiana (Borradaile, 1900), holotype
female (18.9 × 13.5 mm) (SMF 4281), Kadamian River, Sabah.
A, overall dorsal view of habitus; B, frontal view of carapace;C, ventral view showing abdomen, third maxillipeds and chelae.

Fig. 6. Arachnothelphusa kadamaiana (Borradaile, 1900), male (20.1 × 14.9 mm) (SMF 4282), Kadamian River, Sabah. A, right side ofcarapace; B, male abdominal somite 6 and telson; C, ventral view of left G1; D, ventral view of terminal segment of left G1; E, dorsal
view of terminal segment of left G1; F, left G2; G, distal segment of left G2. Setae denuded for all structures. Scale bars: 3.0 mm (A);
1.0 mm (B); 0.5 mm (C, F); 0.25 mm (D, E, G).

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Other true freshwater crabs from the Old World showing
similar phytotelmic behaviour are Potamonautes raybouldi
Cumberlidge & Vannini, 2004 (Potamonautidae) in
Tanzania, East Africa; Globonautes macropus (Rathbun,
1898) (Potamonautidae) in Liberia, West Africa; Malagasya
goodmani (Cumberlidge, Boyko & Harvey, 2002)(Potamonautidae) in Madagascar; and Perbrinckia scansor(Ng, 1995) (Gecarcinucidae) in Sri Lanka (Ng, 1995;
Cumberlidge & Sachs, 1991; Bayliss, 2002; Cumberlidge etal., 2002, 2005; Cumberlidge & Vannini, 2004; Bahir et al.,2008). In Sarawak, the sesarmid crab Scandarma splendidum
has been reported climbing forest trees at night in western
Sarawak (Naruse & Ng, 2007), but it is not known to be aphytotelm crab.

giving us permission to study the freshwater crabs found in
the nearby forests. The first author benefited from a visiting
fellowship from the Lee Kong Chian National History
Museum, National University of Singapore. Michael Türkay(SMF) kindly hosted the third author during his visit to
examine the type of A. kadamaiana. The first two authors
thank Andrew Alek Tuen for administrative support andIndraneil Das for important discussions on many aspects
related to this study. Thanks are due to two reviewers for theirconstructive comments. Research Permit No. 282/2014 was
granted by the Sarawak Forest Department. The study was
supported by The Mohamed bin Zayed Species Conservation
Fund – Project No. 13054042.
LITERATURE CITED

Conservation status. Considering unique habitat andprobable limited distribution, A. merarapensis will easilybe threatened by human activities, especially logging. The
type locality is not protected but the forest is currently safe
because it is part of the Merarap Hot Spring Resort. However,the forest in the vicinity is part of a concession allocated tologging companies and the future of the species is therefore
uncertain. As such, we recommend that A. merarapensis
should at least be listed as “Vulnerable” in the IUCN Red
List of Threatened Species (cf. Cumberlidge et al., 2009).

Bahir MM, Ng PKL, Crandall K, Pethiyagoda R & Cumberlidge N(2008) Perbrinckia scansor. The IUCN Red List of Threatened
Species. Version 2014.2. www.iucnredlist.org. (Accessed 14
November 2014).
Bayliss J (2002) The East Usumbara tree-hole crab (Brachyura:
Potamoidea: Potamonautidae) – a striking example of crustacean
adaptation in closed canopy forest, Tanzania. African Journalof Ecology, 40: 26–34.
Borradaile LA (1900) On a small collection of decapod crustaceansfrom freshwater in North Borneo. Proceedings of the ZoologicalSociety of London, 1900: 93–95.
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, Frankfurt,
526: 1–338, pls. 1–58.
Colosi G (1920) I Potamonidi del R. Museo Zoologico di Torino.
Bollettino dei Musei di Zoologia ed Anatomia comparata della
Regia, Università di Torino, 35(734): 1–39.
Cumberlidge N, Boyko CB & Harvey AW (2002) A new genus andspecies of freshwater crab (Decapoda: Crustacea: Potamoidea)from northern Madagascar, and a second new species associatedwith Pandanus leaf axils. Journal of Natural History, 36(1):65–77.
Cumberlidge N, Fenolio DB, Walvoord ME & Stout J (2005)
Tree-climbing crabs (Potamonautidae and Sesarmidae) fromphytotelmic microhabitats in rainforest canopy in Madagascar.
Journal of Crustacean Biology, 25: 302–308.
Cumberlidge N & Sachs R (1991) Ecology, distribution, and growth
in Globonautes macropus (Rathbun, 1898), a tree-living freshwater crab from the rain forests of Liberia (Parathelphusoidea,
Gecarcinucidae). Crustaceana, 61(1): 55–68.
Cumberlidge N & Vannini M (2004) Ecology and taxonomy
of a tree-living freshwater crab (Brachyura: Potamoidea:
Potamonautidae) from Kenya and Tanzania, East Africa. Journalof Natural History, 38: 681–693.
Cumberlidge N, Ng PKL, Yeo DCJ, Magalhães C, Campos
MR, Alvarez F, Naruse T, Daniels SR, Esser LJ, Attipoe
FYK, Clotilde-Ba F-L, Darwall W, Mcivor A, Collen B &Ram M (2009) Freshwater crabs and the biodiversity crisis:
importance, threats, status, and conservation challenges. Biology
Conservation, 142: 1665–1673.
Grinang, J & Ng PKL (2014) Taxonomy of the freshwater
crab Coccusa cristicervix Ng & Grinang, 2004 (Decapoda:
Brachyura: Gecarcinucidae), with description of a new species
Thelphusula from Sarawak, Malaysia, Borneo. Raffles Bulletinof Zoology, 62: 389–395.
Lanchester WF (1900) On some malacostracous crustaceans from
Malaysia in the collection of the Sarawak Museum. Annals andMagazine of Natural History, 7(6): 249–265, pl. 12.

Key to the species of Arachnothelphusa
1. Carapace surface covered with rough granules; external orbital
angle separated from epibranchial tooth by deep, broad cleft..
..................................................................................................2– Carapace surface smooth, slightly or distinctly rugose; external
orbital angle separated from epibranchial tooth by narrow cleft,teeth sometimes almost confluent............................................3
2. External orbital angle separated from epibranchial tooth bywide, deep cleft; terminal segment of G1 long, about half ofthe length of subterminal segment; distal segment of G2 short,about a quarter length of basal segment (Sarawak)..................
......................................................................... A. merarapensis– External orbital tooth separated from epibranchial tooth bydeep U-shaped cleft; terminal segment of G1 distinctly curved
upwards, short, a quarter length of subterminal segment; distalsegment of G2 very short, about 0.1 times length of basal
segment (Sabah).......................................................A. terrapes
3 Carapace smooth, strongly inflated; branchial regions swollen;outer margin of external orbital angle slightly convex; G1 not
known; (Sabah)............................................. A. rhadamanthysi– Carapace slightly to distinctly rugose; surface and branchialregions convex but not strongly inflated; outer margins ofexternal orbital angle straight or slightly concave..................4
4. Outer margins of external orbital angle smooth; epibranchial
tooth well developed, sharp; terminal segment of G1 short,
one-third length of subterminal segment (central Borneo).......
.............................................................................. A. melanippe– Outer margins of external orbital angle serrate; epibranchial
tooth very low, blunt, almost confluent with external orbital
angle; G1 very slender; terminal segment of G1 short, a quarterof length of subterminal segment (Sabah)....... A. kadamaiana

ACKNOWLEDGEMENTS
We thank Alfred Padan and his family for his kind hospitality
during our visit to the Merarap Hot Spring Resort, and for459

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Ng PKL & Álvarez F (2000) Villalobosus, a replacement namefor Stygothelphusa Alvarez and Villalobos, 1991 (Crustacea:
Brachyura: Pseudothelphusidae), preoccupied by Stygothelphusa
Ng, 1989 (Crustacea: Brachyura: Gecarcinucidae). Raffles
Bulletin of Zoology, 48: 337.
Ng PKL & Goh R (1987) Cavernicolous freshwater crabs (Crustacea,
Decapoda, Brachyura) from Sabah, Borneo. Stygologia, 3(4):313–330.
Ng PKL & Grinang J (2014) A new species of troglobitic crab of
the genus Stygothelphusa Ng, 1989, from Sarawak, Malaysia(Crustacea: Decapoda: Brachyura: Gecarcinudidae). Zootaxa,
3774(1): 90–96.
Rathbun MJ (1898) Descriptions of three species of fresh-water
crabs of the genus Potamon. Proceedings of the BiologicalSociety of Washington, 12: 27–30.
Rathbun MJ (1905) Les crabes d’eau douce (Potamonidae).
Nouvelles Archives du Muséum d’Histoire Naturelle, Paris,(4)7: 159–323, pls. 13–22.
Tan SH & Ng PKL (1998) Revision of the genus Thelphusula(Decapoda: Brahcyura: Gecarcinucidae) with description ofa new genus and three new species. Journal of Crustacean
Biology, 18(4): 808–822.

Man JG De (1899) Zoological Results of the Dutch Expedition
to Central Borneo. The Crustacea. Part II, Brachyura. Notes
Leyden Museum, 21: 53–144, pls. 5–12.
Naruse T & Ng PKL (2007) Scandarma splendidum, a new speciesof tree-climbing crab (Decapoda: Brachyura: Sesarmidae)
from Sarawak, Malaysia, Borneo. Raffles Bulletin of Zoology,
55(2): 337–341.
Ng PKL (1988) The freshwater crabs of Peninsular Malaysia andSingapore. Department of Zoology, National University ofSingapore, Shinglee Press, Singapore, i–viii, 1–156 pp., figs.
1–63, 4 colour pls.
Ng PKL (1989) The identity of the cavernicolous freshwater crab
Potamon (Thelphusa) bidiense Lanchester, 1900 (Crustacea:
Decapoda: Brachyura: Gecarcinucidae) from Sarawak, Borneo,
with description of a new genus. Raffles Bulletin of Zoology,
37(1&2): 63–72.
Ng PKL (1991) Bornean freshwater crabs of the genus
Arachnothelphusa gen. nov. (Crustacea: Decapoda: Brachyura:
Gecarcinucidae). Zoologische Mededelingen, 65: 1–12.
Ng PKL (1995) Ceylonthelphusa scansor, a new species of treeclimbing crab from Sinharaja Forest in Sri Lanka (Crustacea:
Decapoda: Brachyura: Parathelphusidae). Journal of South
Asian Natural History, 1(2): 175–184.
Ng PKL (2013) Stygothelphusa cranbrooki, a new species of cavecrab from Gua Sireh, Sarawak, Malaysia (Crustacea: Decapoda:
Brachyura: Gecarcinudidae). Raffles Bulletin of Zoology,
Supplement 29: 91–97.

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