THE RAFFLES BULLETIN OF ZOOLOGY 2013
THE RAFFLES BULLETIN OF ZOOLOGY 2013 Supplement No. 29: 91–97
http://zoobank.org/urn:lsid:zoobank.org:pub:0AD662A2-851E-49A6-B8D4-4297878EE4A1
Date of Publication: 30 Nov.2013© National University of Singapore

STYGOTHELPHUSA CRANBROOKI, A NEW SPECIES OF CAVE CRAB
FROM GUA SIREH, SARAWAK, MALAYSIA(CRUSTACEA: DECAPODA: BRACHYURA: GECARCINUCIDAE)Peter K. L. Ng
Raffles Museum of Biodiversity Research, Faculty of Science, National University of Singapore
14 Science Drive 4, 117543 Singapore
Email: dbsngkl@nus.edu.sg

ABSTRACT. — A new species of freshwater cave crab of the genus Stygothelphusa Ng, 1989 (Gecarcinucidae)
is described from Gua Sireh in Sarawak, Malaysia. The species is allied to S. bidiensis (Lanchester, 1900) andS. nobilii (Colosi, 1920) from other parts of Sarawak, but differs by its carapace physiognomy, pronounced
epibranchial tooth, proportionately much longer ambulatory legs and distinctive male first gonopod structure.
KEY WORDS. — Crustacea, Brachyura, Gecarcinucidae, Stygothelphusa, new species, cave, Sarawak,
Malaysia

INTRODUCTIONTAXONOMY

Ng (1989) established a new genus, Stygothelphusa, for
Potamon (Thelphusa) bidiense Lanchester, 1900, a species
orginally described from the Bidi cave system in Bau,
Sarawak. A second poorly known species, Parathelphusa(Liothelphusa) nobilii Colosi, 1920, from Sarawak, originallyplaced in the Bornean genus Arachnothelphusa Ng, 1991,
was later referred to Stygothelphusa by Ng & Álvarez (2000)(see also Ng, 2004; Ng & Yeo, 2007; Ng et al., 2008).

Family Gecarcinucidae Rathbun, 1904
Stygothelphusa Ng, 1989
Type species. — Potamon (Thelphusa) bidiense Lanchester, 1900,
by original designation; gender feminine.

Remarks. — Stygothelphusa was established for the typespecies by Ng (1989) because of its squarish carapace, very
long ambulatory legs, distinctive G1 structure which has
the terminal segment less than a quarter of the total length,
and a G2 which has an elongated distal segment which is
as long as or equal to the basal segment (Ng, 1989, 2004).

In the 1990s, the author collected numerous specimensfrom a cave system about 60 km east of the from the Bau
system in Kuching that superficially resembled S. bidiensis
and S. nobilii but differed markedly in having a pronounced
epibranchial tooth, relatively longer ambulatory legs and adifferently shaped male first gonopod. It is here described
as a new species, S. cranbrooki.

The taxonomy of the type species, Stygothelphusa bidiensis(Lanchester, 1900) has been treated at length by Ng (1989)
and there is no need to elaborate. The type specimen is nolonger extant, and a neotype male from the Bidi caves was
selected by Ng (1989: 70).

The abbreviations G1 and G2 are used for the male firstand second gonopods, respectively. The terminology usedfollows Ng (1988). Measurements provided (in millimetres)
are of the carapace width and length, respectively. Specimens
examined are deposited in the Sarawak Museum, Kuching(SM); Museum of Zoology, University of Turin (MZUT);
and the Zoological Reference Collection of the Raffles
Museum of Biodiversity Research, National University ofSingapore (ZRC).

Parathelphusa (Liothelphusa) nobilii Colosi, 1920, is apoorly known species. It was originally reported as “Potamon(Geothelphusa) kenepai De Man, 1899” by Nobili (1903:
15–16) from two male and one female specimens collectedfrom 2500 feet on Gunung Saribau (= Sari-Baw) by
Robert Shelford in 1902. This location is uncertain but isunlikely to be what is today called Saribas near Sri Aman(ca. 1°10'N, 111°38'E) to the east of Serian. Interestingly,
Shelford (1905: 210) described a new frog from this location
which he reported as “Mount Saribaw, Samarahan River,91

Ng: New species of Stygothelphusa from Sarawak, Malaysia
Sarawak”. The Samarahan area is closer to the Serian and
Bau areas, and also has large sections of limestone, and islikely Shelford’s crab specimens came from this area. Colosi(1920: 25) commented that Nobili had originally believed
his specimens belonged to a new species and he gave aprovisional name, “Potamon saribanensis”, which was neverpublished. Colosi (1920) recognised the Saribau specimens as
new, and named them Parathelphusa (Liothelphusa) nobilii.
Although both Nobili (1903) and Colosi (1920) described thespecies, no figures have ever been provided of the taxon andits gonopods are not known, so its precise generic affinities
could not be ascertained with any certainty. Bott (1970: 59)
synonymised Colosi’s species with Potamon (Geothelphusa)
hendersonianum De Man, 1899 (from central Kalimantan),
without comment and referred it to Thelphsula Bott, 1969.
Ng (1995: 196) discussed the problems with Bott’s (1970)classification of Potamon kenepai (originally described from
Mount Kenepa in central Kalimantan) (which he referred
to Adeleana Bott, 1969) and P. hendersonianum (which heplaced in Thelphusula Bott, 1969). Ng (1995) argued thatboth species were closely related, and transferred them toa new genus, Bakousa Ng, 1995 (type species Bakousa
sarawakensis Ng, 1995).

not associated with caves and have pigmentated carapaces
and legs (De Man, 1899; Ng, 1995), while Stygothelphusa isalways associated with caves and has pale body colouration.
The G1 and G2 differences are not substantial, and while theG1 terminal segment of Stygothelphusa species is slightly
stouter (for S. bidiensis at least), and the G2 basal segment isrelatively shorter compared to the long distal segment, whatstands out are the much longer pereopods of Stygothelphusa.
Another key character to separate Stygothelphusa from
Bakousa is the form of the anterior male thoracic sternum(sternites 1–4). In Stygothelphusa, the anterior thoracic
sternum is proportionately narrower transversely with thesternoabdominal cavity reaching to the level of the junction
between sternites 2 and 3, on an imaginary line connecting
the anterior margins of the coxae of the chelipeds (Figs.
3C, 4A). In Bakousa, the anterior male thoracic sternum is
proportionately wider transversely, and the sternoabdominal
cavity only reaches to the junction between sternites 3 and4, on an imaginary line connecting the median points of thecoxae of the chelipeds (Ng, 1995: Fig. 12C).
At the author’s request, Oliver Chia examined andphotographed the three type specimens of Parathelphusa(Liothelphusa) nobilii in MZUT and it is clearly a species of
Stygothelphusa (Fig. 2). The form of the male anterior thoracic

Ng (1995: 197) noted that Bakousa was related to
Stygothelphusa, but noted that the former had much shorterchelipeds and ambulatory legs, a proportionately longer
ischium of the third maxillipeds, a relatively stouter G1 and ashorter G2 basal segment. All Bakousa species are free-living,

Fig. 2. Stygothelphusa nobilii (Colosi, 1920). Overall habitus. A,
lectotype male (15.0 × 13.0) (MZUT Cr 1308), Saribau, Sarawak;
B, paralectotype female (21.0 × 18.0 mm) (MZUT Cr 1308);
C, paralectotype male (17.0 × 14.7 mm) (MZUT Cr 1308). All
specimens from Saribau, Sarawak.

Fig. 1. Stygothelphusa bidiensis (Lanchester, 1900). A, C, male
(15.4 × 13.7 mm) (ZRC 1998.539); B, D, male (12.9 × 11.9 mm)(ZRC 2013.1624). A, B, overall habitus; C, D, dorsal views ofcarapaces. Specimens from Fairy Cave, Bau, Sarawak.

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Comparative material. — Stygothelphusa bidiensis (Lanchester,1900): 1 neotype male (16.4 × 14.5 mm) (SM Cru 1986.17), Bidi
Caves, bau, Sarawak, coll. C. J. Brooks, Jun.1903; 2 males, 3
females (largest 16.9 × 14.8 mm), 2 juveniles (SM Cru 1986.18–25),
Bidi Caves, Bau, Sarawak, coll. C. J. Brooks, Jun.1903; 2 males(larger 12.9 × 11.9 mm), 2 females (larger 17.2 × 15.7 mm) (ZRC
2013.1624), Gua Kapo (= Fairy Cave), Bau Caves, Sarawak,
1°22.91'N, 110°7.08'E, coll. P. K. L. Ng & H. H. Tan, 28 Oct.1997;
3 males (largest 11.9 × 9.8 mm), 6 females (largest 16.3 × 13.7mm) (ZRC 2013.1623), Gua Kapo (= Fairy Cave), Bau Caves,
Sarawak, 1°22.91'N, 110°7.08'E, coll. H. H. Tan & P. K. L. Ng,
28 Oct.1997; 4 males (largest 15.4 × 13.7 mm) (ZRC 1998.539),
Gua Kapo (= Fairy Cave), Bau Caves, Sarawak, 1°22.91'N,
110°7.08'E, coll. students, 23 Jun.1998; 1 female (15.9 × 13.6mm) (ZRC 1989.3038), Gua Pitas, Bau Caves, Sarawak, coll. S.
Yussof, 1987; 2 males (larger 15.0 × 13.0 mm), 2 females (larger
17.0 × 14.9 mm) (ZRC 1984.7543–7546), Borneo, no other data.
Stygothelphusa nobilii (Colosi, 1920): lectotype male (15.0 × 13.0mm) (here designated), 1 paralectotype male (17.0 × 14.7 mm), 1
female (20.8 × 18.1 mm) (MZUT Cr 1308), 2500 feet asl, Gunung
Saribau, Sarawak, coll. R. Shelford, 1902.

sternum and third maxillipeds agree with Stygothelphusa as
discussed above. All three specimens carry the catalogue
number MZUT Cr 1308. Bott (1970: 59) listed a “lectotype”
male specimen in the museum with the number 1570 but he
did not give measurements so we do not know which one
it was. Bott (1970: 59) also listed a specimen measuring21.0 by 18.0 mm but its sex and type status was not stated.
Colosi (1920: 26) had noted that the type specimens of
Parathelphusa (Liothelphusa) nobilii had been labelled “n.
1570 Potamon saribanensis Nob. Tipi : Sarawak, Monte
Sariban, R. Shelford 1902”, and Bott probably cited Nobili’s
number. Whether Bott actually examined Colosi’s types (orphotographs) is not known.
Nobili (1903: 15, 16) noted that he only had one complete
male measuring 15.0 by 12.3 mm, and another larger
incomplete male 17.0 mm in carapace width. The female was
not measured. The most complete male photographed is 15.0
× 13.0 mm (differing slightly from Nobili’s measurements forthe length) (Fig. 2A). The larger male has lost its chelipeds
and several of its legs is 17.0 × 14.7 mm (Fig. 2C). The
female specimen is 20.8 × 18.1 mm (Fig. 2B), which agrees
well with the measurement of 21.0 × 18.0 mm given in Bott
(1970: 59), suggesting he measured the largest specimen in theseries. The most complete male specimen (15.0 × 13.0 mm)(Fig. 2A) is here designated the lectotype of Parathelphusa(Liothelphusa) nobilii Colosi, 1920.

Diagnosis. — Carapace quadrate, broader than long; dorsal
surfaces gently rugose (especially along margins) to smooth(Fig. 3A, B); striae on anterolateral regions strong (Fig. 3B);
branchial regions gently inflated dorsally and laterally, lateral
margins appearing gently convex from dorsal view (Fig. 3B,D), gently convex from frontal view (Fig. 3D); epibranchial
tooth spiniform, distinct, separated from external orbital
angle by clear cleft (Fig. 3A, B); ambulatory legs very long

The carapace of Stygothelphusa nobilii is superficiallyvery similar to S. bidiensis, but can easily be separated byits proportionately shorter ambulatory legs (see discussion
for S. cranbrooki later). Its G1 structure is not known. As
discussed earlier, the precise type locality of Stygothelphusa
nobilii is uncertain.
The nomenclatural problems associated with a juniorhomonym, Stygothelphusa Álvarez & Villalobos, 1991,
established for a cave species from Mexico, has been
discussed at length by Ng & Álvarez (2000) and Ng &Low (2010).

Stygothelphusa cranbrooki sp. nov.
(Figs. 3, 4, 5C, D, F, 6E–H)
Material examined. — Holotype: male (15.0 × 13.6 mm) (ZRC
2013.1621), Gua Sireh, near Kampung Bantang, Sarawak,
1°10.71'N, 110°28.09'E, coll. P. K. L. Ng & H. H. Tan, 30 Oct.1997.
Paratypes: 1 female (brooding young crabs) (15.9 × 13.7 mm)(ZRC 2013.1620), coll. P. K. L. Ng & H. H. Tan, 30 Oct.1997;
5 males (largest 19.4 × 16.4 mm), 6 females (largest 16.6 × 14.5mm) (ZRC 2013.1622), coll. students 30 Oct.1997; 1 male (13.8
× 12.4 mm) (ZRC 2013.1619), coll. P. K. L. Ng, 23 Jun.1998; 6
males (largest 15.4 × 13.9 mm), 6 females (largest 17.2 × 15.0mm) (ZRC 1998.540), coll. students, 23 Jun.1998; 1 male (10.8
× 9.4 mm), 3 females (largest 16.1 × 14.5 mm) (ZRC 1998.541),
coll. P. K. L. Ng, 25 Jun.1998; 5 males (largest 16.8 × 14.9 mm),
5 females (16.6 × 14.9 mm) (ZRC 1999.690), coll. P. K. L. Ng, 10
Jun.1999. All specimens from Gua Sireh, near Kampung Bantang,
Sarawak, 1°10.71'N, 110°28.09'E.

Fig. 3. Stygothelphusa cranbrooki sp. nov. Holotype male (15.0
× 13.6 mm) (ZRC 2013.1621), Gua Sireh, Sarawak. A, overall
habitus; B, dorsal view of carapace; C, anterior thoracic sternum
and third maxillipeds; D, frontal view of carapace.

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Ng: New species of Stygothelphusa from Sarawak, Malaysia(Fig. 3A); male sternoabdominal cavity reaching to level ofjunction between thoracic sternites 2 and 3, on imaginaryline connecting anterior margins of coxae of chelipeds(Figs. 3C, 4A); male abdomen T-shaped (Fig. 4); G1 curved
outwards; terminal segment gently curved upwards, ca. 0.2
times subterminal segment (Fig. 6E-G); G2 distal segment
flagelliform, subequal in length to basal segment (Fig. 6H).

spines; dactylus gently curved, with 2 rows of spines on
dorsal and ventral margins (Figs. 3A, 5F).
Male abdomen T-shaped; telson elongate, lateral margins
almost straight, tip rounded, as long as somite 6 (Fig. 4A);
somites 3–6 progressivley narrower, trapezoidal (Fig. 4B);
somite 3 widest, covering most of thoracic sternite 8 (Fig.
4C); somites 1 and 2 longitudinally narrow, reaching tocoxae of last pair of ambulatory legs (Fig. 4C).

Description of holotype male. — Carapace quadrate, broader
than long; regions distinct; dorsal surfaces gently rugose
to smooth; antero- and posterolateral regions lined with
distinct oblique striae; cervical grooves broad but shallow;
pterygostomial, suborbital and sub-branchial regions rugose
to granulose (Fig. 3). Branchial regions gently inflated
dorsally and laterally (Fig. 3A, B, D). Frontal margin distinct,
level with tip of external orbital tooth, gently sinuous,
median part gently deflexed downwards, margin forming
poorly defined median pseudo-frontal triangle (Fig. 3B, D).
Anterolateral margin gently convex, not clearly demarcatedfrom posterolateral margin; epibranchial tooth distinct, sharp,
separated from external orbital angle by distinct cleft; external
orbital angle acutely triangular, outer margin ca. 2 times
length of inner margin, lined with small granules; striae
on anterolateral regions distinct, strong; epigastric cristae
strong, distinctly raised, rugose but not sharp, separated
by deep median Y-shaped groove, anterior of postorbitalcristae; postorbital cristae relatively low, indistinct but sharp,
separated from epigastric cristae y shallow cervical groove(Fig. 3A, B, D). Posterior margin of carapace gently sinuous(Fig. 3A, B). Epistome wide; posterior margin with distinct
median triangular lobe with rounded tip, lateral margins
gently sinuous (Fig. 3D). Eye distinct, filling most of orbit;
cornea large, fully pigmented (Fig. 3B, D). Antennular fossa
transversely narrow, rectangular in shape; flagellum folding
transversely (Fig. 3D). Third maxilliped quadrate; ischium
quadrate, with shallow oblique median sulcus; merus squarish
with slightly auriculiform anteroexternal margin; exopod
reaches beyond distal edge of ischium, flagellum distinct,
longer than width of merus (Fig. 3C). Thoracic sternum
smooth, relatively narrow transversely, sternites 1–4 fused,without distinct median sutures visible; sternoabdominal
cavity reaching to the level of the junction between sternites
2 and 3, on an imaginary line connecting the anterior marginsof the coxae of the chelipeds (Figs. 3C, 4A).

G1 with proximal part stout, straight, distal half curved
outwards; terminal segment tapering towards subtruncate
tip, gently curved upwards (towards buccal cavity in situ),
ca. 0.2 times subterminal segment, distal part covered withnumerous very small scale-like spinules, rest of margin withlong and short setae (Fig. 6E–G). G2 much longer than G1;
basal segment long, distal segment flagelliform, subequal inlength to basal segment (Fig. 6H).
Variation. — Adult female specimens are similar to the males
in all non-sexual characters, except that their chelipeds are
more symmetrical. Female abdomens are wide, rounded, andcompletely cover the thoracic sternum. The epibranchial tooth

Chelipeds elongated; left larger; outer surfaces and margins
distinctly rugose, without sharp spines or spinules; distal edgeof dorsal margin of elongate merus with spine; carpus with
low inner distal spine, posterior margins lined with small
granules; outer surface of palm gently (Figs. 3A, 5C, D);major chela with fingers shorter than palm, cutting edge ofpollex with 2 teeth and numerous denticles, cutting edgeof dactylus with submedian tooth and denticles (Fig. 5C);
minor chela with fingers longer than palm, cutting edges offingers with numerous denticles (Figs. 5D).
Fig. 4. Stygothelphusa cranbrooki sp. nov. Holotype male (15.0
× 13.6 mm) (ZRC 2013.1621), Gua Sireh, Sarawak. A, anterior
thoracic sternum, abdominal somite 6 and telson; B, thoracic
sternum and abdominal somites 3–5; C, posterior thoracic sternum
and abdominal somites 1–3.

Ambulatory legs very long, slender; second pair longest;merus unarmed, dorsal margin gently serrated but notspiniform, with low subdistal spine; outer surfaces in firstto third pairs rugose; propodus with row of small ventral

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Etymology. — Gua Sireh is a significant archaeological site,and may have been occupied for some 20,000 years. It is
an important site for pottery and other artefacts, and has
ancient charcoal drawings on some of the walls (see Datan& Bellwood, 1991; Doherty et al., 2000). In recent years,
Gua Serih (and the adjacent areas) has become commercially
important for the harvest of bird’s nests from swiftlets. The
naming of this species after Gathorne Gathorne-Hardy, theEarl of Cranbrook (better to known to scholars as Lord
Medway), a renowned archaeologist and expert on cave
swiftlets, is therefore all the more appropriate. I therefore
take great pleasure in naming this species after an old friend
who has contributed so much to Southeast Asian biodiversity
and archaeology!

is usually distinct although in some specimens, it appears to
be partially worn down; although in all the specimens, thestriae on the anterolateral regions are always stronger andmore prominent compared to S. bidiensis.
Colour. — In life, S. cranbrooki is white to light yellow
overall; although the cornea of the eyes are pigmented black.
Habitat. — Stygothephusa cranbrooki was only found in
the dark zone of the cave and was not observed near theentrance. The main opening to the cave is some 50 m above
the ground and opens into a complex cave system that leads
downwards into subterranean rivers. The crabs were collectedin moist areas under rocks in the dark zone, and were very
common in places where there was water and piles of rubble.

Remarks. — Stygothelphusa cranbrooki can easily beseparated from S. bidiensis in having a well developed sharp
epibranchial tooth clearly separated from the anterolateral
margin by a deep cleft (Fig. 3A, B) (versus poorly developed,
sometines not discernible, not separated from anterolateral
margin by cleft, Fig. 1), the striae on the anterolateral regions
are always relatively stonger and more pronounced (Fig.
3B versus Fig. 1C, D); the branchial regions are distinctly
more swollen, with the surfaces appearing gently convex
from frontal view, and the lateral margins of the carapace
appearing squarish (Fig. 3A, B) (versus less swollen, withthe surfaces appearing distinctly convex from frontal view,
resulting in the lateral margins of the carapace appearing
more rounded in shape, Figs. 1, 5A), and the G1 terminal
segment is proportionately more elongate and gently curved
upwards (Fig. 6E–G) (versus relatively shorter, straighter andconical, Fig. 6A–C). In addition, most of the adult specimensof S. cranbrooki have proportionately longer ambulatory
legs (especially the merus) compared to S. bidiensis (Fig.
5F versus Fig. 5E), although this character is not always
reliable, with some large female specimens of S. bidiensis
also having relatively longer legs.

The behaviour and habits of S. cranbrooki are the same as
those described for S. bidiensis (see Ng, 1989; Ng & Yussof,
1990). While it has some cave-adapted characters like very
long pereopods and a general loss of body pigmentation,
the eyes are not degenerated and the cornea retains all of its
pigmentation. While it cannot be regarded as a troglobite (cf.
Holthuis, 1986; Guinot, 1988), it has, however, also neverbeen found outside caves. In Gua Sireh, S. cranbrooki has
been found in good numbers over 100 m deep inside thecave, in complete darkness.

The carapace physignomy of S. cranbrooki is similar to
S. nobilii in that their branchial regions are relatively lessswollen (Figs. 2, 3A, B, D) than those of S. bidiensis (Figs.
1, 5A). However, the condition of the epibranchial spine of S.
nobilii more closely resembles that of S. bidiensis, being low
and not as clearly separated from the rest of the anterolateral
margin (Figs. 1, 2) compared to S. cranbrooki (Fig. 3A, B).
The ambulatory legs of S. nobilii are also distinctly shorter
than those of S. bidiensis and S. cranbrooki. When folded
against the sides of the carapace, the last ambulatory merusof S. nobilii does not reach the frontal margin (Fig. 2). In
S. bidiensis and S. cranbrooki, the last ambulatory merus
reaches or extends beyond the frontal margin of the carapace(Figs. 1A, B, 3A).
The conservation status of S. cranbrooki has been discussed
under S. bidiensis sensu lato (see Ng & Yeo, 2007;
Cumberlidge et al., 2009). The species is listed as vulnerable
but the status of both species will need to be reassessed inview of the revised taxonomy here. Stygothelphusa bidiensis
is now restricted the limestone formations in Bau, while S.
cranbrooki is known only from the Gua Sireh area.

Fig. 5. A, B, E, Stygothelphusa bidiensis (Lanchester, 1900), male
(15.4 × 13.7 mm) (ZRC 1998.539), Fairy Cave, Bau, Sarawak; C,
D, F, Stygothelphusa cranbrooki sp. nov., holotype male (15.0 ×
13.6 mm) (ZRC 2013.1621), Gua Sireh, Sarawak. A, frontal viewof carapace; B, outer view of right major chela; C, outer view ofleft major chela; D, outer view of right minor chela; E, F, rightfourth ambulatory legs.

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Ng: New species of Stygothelphusa from Sarawak, Malaysia
ACKNOWLEDGEMENTS

LITERATURE CITEDThis paper is a contribution to Supplement No. 29 of the
Raffles Bulletin of Zoology, marking the eightieth birthdayof the Earl of Cranbrook (V). Thanks are due to Charles
Leh, chief zoologist at the Sarawak Museum, for hostingand helping so many of the author’s research sojourns inSarawak over the years; and for originally suggesting wesample at Gua Sireh. Thanks are also due to Oliver Chia whokindly helped me check and photograph the type specimens
in MZUT during his visit there.

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Fig. 6. A–D, Stygothelphusa bidiensis (Lanchester, 1900), male (12.9 × 11.9 mm) (ZRC 2013.1624), Fairy Cave, Bau, Sarawak; E–H,
Stygothelphusa cranbrooki sp. nov., holotype male (15.0 × 13.6 mm) (ZRC 2013.1621), Gua Sireh, Sarawak. A, E, left G1, ventral views;
B, F, terminal segments of left G1s, ventral views; C, G, terminal segments of left G1s, dorsal views; D, H, left G2s. Scales bars = 0.5
mm (A, D, E, H), 0.1 mm (B, C, F, G).

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