THE RAFFLES BULLETIN OF ZOOLOGY 2007
THE RAFFLES BULLETIN OF ZOOLOGY 2007 Supplement No. 16: 79–94
Date of Publication: 31 Dec.2007© National University of Singapore

LEUCOSIID CRABS FROM PANGLAO, PHILIPPINES, WITH DESCRIPTIONS OF THREENEW SPECIES (CRUSTACEA: DECAPODA: BRACHYURA)
Bella S. Galil
National Institute of Oceanography, Israel Oceanographic and Limnological Research
P.O.B. 8030, Haifa 31080, Israel
Email: bella@ocean.org.il

Peter K. L. Ng
Department of Biological Sciences, National University of Singapore,
14 Science Drive 4, Singapore 117543, Republic of Singapore
Email: dbsngkl@nus.edu.sg

ABSTRACT. – Thirty-eight species of leucosiid crabs are reported from Panglao in Bohol, the Central
Philippines. Of these, three are new to science: Alox bothros, A. chaunos, and Urnalana cristata, while five
constitute new records for the Philippines: Leucosia rubripalma Galil, 2003, Myra tumidospina Galil, 2001,
Urnalana elata (A. Milne-Edwards, 1874), U. pulchella (Bell, 1855) and U. whitei (Bell, 1855). The new
species are described and illustrated, and their affinities with allied taxa discussed Tokoyo triloba Komatsu,
Manual & Takeda, 2005, is also synonymised with T. eburnea (Alcock, 1896)
KEY WORDS. – Crustacea, Decapoda, Brachyura, Leucosiidae, Alox, Urnalana, new species, Philippines.

INTRODUCTION

Panglao and around Balicasag Island was obtained mostly
by trawls and dredges, with specimens also coming fromcoral-brushings (from rubble collected by divers), diving aswell as hand-collections from intertidal areas. A good part ofthe material from Balicasag Island, however, was collectedby local shell fishermen using tangle nets set to depths of
500 m (see also McLay & Ng, 2005). These collections andexpeditions resulted in extensive series of specimens ofleucosiid crabs. On the basis of the studied material, 38
species have been identified to date. Of these, three are new
to science: Alox bothros, A. chaunos, and Urnalana cristata,and five constitute new records for the Philippines: Leucosia
rubripalma Galil, 2003, Myra tumidospina Galil, 2001,
Urnalana elata (A. Milne-Edwards, 1874), U. pulchella (Bell,1855), and U. whitei (Bell, 1855).

The Philippines have featured prominently in our growing
knowledge of the Leucosiidae of the Indo-West Pacific. The
leucosiid crab material from the U.S. Exploring Expedition(1838–1842), Challenger Expedition (1873–1876), Siboga
Expedition (1899–1900), Albatross Expedition (1907–1910),
and the three MUSORSTOM expeditions conducted under theaegis of the Muséum national d’Histoire naturelle, Paris(1976, 1980, 1985) have been studied in part or whole andyielded numerous new and rare species (Bell, 1855a–c; Miers,
1886; Ihle, 1918; Serène & Vadon, 1981; Chen, 1989; Tan,
1996; Tan & Ng, 1995; Galil, 2001a, b, 2003a–c, 2005a, b).
The studies by Philippine carcinologists Roxas (1930) andEstampador (1937, 1959) have also added many new records.
Two recent contributions by Komatsu et al. (2004, 2005) arealso noteworthy for the Philippines.

The present study complements the previous report on theleucosiid crabs from Balicasag Island by Komatsu et al.
(2005) which reports 28 species, two of which were describedas new. Interestingly, the present study and that by Komatsu
et al. (2005) have only 15 species in common. The greaternumber of the genera and species reported in this study canbe explained by the more extensive sampling in more areasand habitats. Together, the two studies highlight the very richcrustacean diversity present in this part of the Philippines. It
must also be emphasised that not all the leucosiid specimensfrom the present series of collections and expeditions haveSince early 2000, the Raffles Museum of Biodiversity
Research (RMBR), Singapore, has been conducting studiesof the crab fauna of Balicasag Island and the nearby Panglao
area in Bohol, the Philippines, with the University of San
Carlos (Cebu, the Philippines). Between 2004 and 2005, theRaffles Museum RMBR joined with the Musèum nationald’Histoire naturelle (Paris, France) and Philippine National
Museum (Manila) to conduct two expeditions to Panglao(including Balicasag Island). The expedition material from79

Galil & Ng: Some Leucosiidae from the Philippines
been studied. We have on hand specimens which in the older
literature would have been identified as “Leucosia anatum(Herbst, 1783)” but as this group of species is now being
studied as part of a larger revision of the genus Leucosia sensu
lato (see Galil, 2003b, c, 2005a, b, 2006a, b), it is excluded
for the present work. In addition, there are still many lots ofvery small leucosiids (notably Nursia and allies,
Cryptocnemus, Ebalia etc.) which have not yet been studied,
and will certainly include more new species and new records.
These are being consolidated for a follow-up study by H.
Komatsu and his collaborators.

prominent, irregularly indented. Longitudinal median ridge
minutely granulate, extending from frontal margin to
granulate cardiac region. Swollen intestinal region, separated
from branchial regions by deep, irregularly granule-lined
grooves. Chelipeds subequal, robust. Cheliped merus trigonalin cross-section; upper surface proximally with cluster of
granules; posterior margin distally bilobed. Palm obscurely
granulate, pitted; fingers twice as long as palm; dactylus
slightly widened distally, upper margin carinate, closely
granulate; lower margin of pollex distally carinate, granulate.
Pereiopods stocky, short; meri with single granulate row on
dorsal margin, two rows on ventral margin; carpi, propodi,
dactyli minutely granulate. Thoracic sternum with transverse
ridges, with 3 granule-lined grooves between them. Male
abdominal sulcus deep, reaching buccal cavity; lateral margin
bearing distinct ridge fitting into suture between abdominal
segments. Abdomen closely covered with flattened granules;
male abdominal segments 1 and 2 slender, horizontal;
segments 3–5 fused; segment 6 large, trapezoid, as long as
laciniate telson. Shaft of G1 angled distad, medially setose.
G2 short, slender, apex scoop-like.

In the present paper, a checklist of the 38 species is presented.
As many species have been previously reported from thePhilippines, short notes are provided for the new records, andcolour descriptions where none was available before; while
the new species are fully described and illustrated.
Holotype specimens examined are deposited in the National
Museum Carcinological Reference Collection (NMCR) of the
National Museum of the Philippines. Other specimens will
be separated at a later date and deposited in the NMCR;
Zoological Reference Collection (ZRC) of the Raffles
Museum of Biodiversity Research, National University ofSingapore; and Musèum national d’Histoire naturelle, Paris(MNHN). Comparative material from the Zoological Museum
Amsterdam (ZMA), Universiteit van Amsterdam, was also
examined. The following abbreviations are used: CL –carapace length, measured along longitudinally across themedian line of the carapace; coll. – collected by; G1 – malefirst pleopod; G2 – male second pleopod.

Colour. – Carapace white, carapacial pits and grooves linedwith orange markings; meral-carpal joint of cheliped bearing
a pair of reddish spots, fingers red-spotted; pereiopodal meri
with orange marbling proximally (Fig. 1A).
Remarks. – Alox bothros, new species, is distinguished fromthe closely related A. somphos Tan & Ng, 1995, in having amore prominent denticle on subhepatic margin, the dorsal
surface of carapace being relatively less pitted, in having anauriculate carapace rim laterally, and in the much stouter formof the G1 (cf. Tan & Ng, 1995). Tan & Ng (1995) did notillustrate the G1 of A. somphos but merely described it as along and slender structure. Together with the male abdomen,
it is figured here (Fig. 5C–G) for comparisons. The distal
part of the G1 of A. somphos is damaged (Fig. 5C, D) but thebroken part was still present and has also been figured together
with the rest of the structure (Fig. 5E).

TAXONOMY
Alox bothros, new species(Figs. 1A, 5A, B)
Material examined. – Holotype, male (CL 9.0 mm) (NMCR), station
S8, Momo Beach, Panglao, 09°36.5'N 123°45.6'E, 28–32 m, cavesin reef wall, 10 Jul.2004.

Etymology. – bothros Greek, “trench, pit”.

Description. – Carapace subpentagonal, 1.6 times as wide as
long. Dorsal surface of carapace prominently and irregularly
sculpted. Front produced, up-curved, margin bilobed, bearing
granulated pit medially. Anterior margin of carapace vertical,
basal antenular segment operculiform, rugose, entirely sealingovate antennular fossa. Antennae folded in orbital hiatus
without gap. Orbits small, rounded, visible in dorsal view;
cornea visible when eye retracted into orbit; outer orbital
margin with 2 sutures. External maxillipeds concealing
trapezoid buccal opening, rugose; endognathal meri visible
in anterior view. Postfrontal region laterally with irregularly
pitted trench, pits surfaced with flattened granules.
Anterolateral margin distinctly rimmed, with lozenge-shaped
indentation medially. Subhepatic margin with large granulate
denticle medially, visible in dorsal view. Lateral margins ofcarapace expanded, anteriorly trilobate, bearing swollen
auriculate carina. Posterior margin bearing triangular denticle
submedially, obscurely bilobate medially. Branchial regions

Distribution. – This species is known only from type location,
Panglao Island.

Alox chaunos, new species(Figs. 1B, 5H, I)
Material examined. – Holotype: male (CL 6.8 mm) (NMCR), station
M7, Momo Beach, Panglao, 09°36.1'N 123°45.2'E, 0–3m, reef
platform with seagrass, 1 Jun.2004. – Paratype: 1 ovigerous female(CL 9.8 mm) (ZRC), Pontod Islet, Panglao, station D4, soft bottom
with sea-grass, 0–2 m, 5 Jun.2004.

Description. – Carapace subpentagonal, about 1.3 as wide as
long. Dorsal surface of carapace closely granulate, pitted,
prominently sculpted. Front produced, up-curved, marginbilobed, swollen. Anterior margin of carapace vertical, basal
antenular segment operculiform, rugose, entirely sealing

80THE RAFFLES BULLETIN OF ZOOLOGY 2007
subtriangular antennular fossa. Antennae folded in orbital
hiatus without gap. Orbits small, rounded, visible in dorsal
view; when retracted, ocular peduncle seals orbit; outer orbital
margin bisutured. External maxillipeds concealing trapezoid
buccal opening, closely set with flattened granules;
endognathal meri visible in anterior view. Postfrontal region
laterally concave. Anterolateral margin indistinctly rimmed,
with shallow indentation medially. Sigmoid, granule-lined
groove stretching from postfrontal ridge to branchial region.
Subhepatic margin with triangular facet medially, visible indorsal view. Lateral margins of carapace expanded, slightly
constricted medially. Posterior margin horizontal
submedially, obscurely bilobate medially. Branchial regions
swollen, irregularly pitted, a pair of nostril-like cavities
submedially on anterior margin. Cardiac region surfaced withflattened granules. Intestinal region well demarcated,
separated from branchial regions by granule-lined groove.
Chelipeds subequal, robust. Cheliped merus trigonal in crosssection; upper surface proximally with cluster of granules;
posterior margin distally bilobed. Palm obscurely granulate,pitted; fingers as long as palm, closely granulate, obscurely
carinate. Pereiopods stocky, short; meri with single granulate
row on dorsal margin, 2 rows on ventral margin; carpi,
propodi, dactyli minutely granulate. Thoracic sternum closely
granulate, horizontally ridged, ridges interspaced with 3
granule-lined grooves. Male abdominal sulcus deep, reaching
buccal cavity. Abdomen closely covered with flattened
granules; male abdominal segments 1 and 2 slender; segments
3–5 fused; segment 6 trapezoid; telson narrow, elongate,
distinctly longer than segment 6. Shaft of G1 almost straight,
tapering, distally setose. G2 short, slender, apex scoop-like.

Alox ornatum (Ihle, 1918)(Fig. 1C)
Oreophorus (Oreophorus) ornatus Ihle, 1918: 214, Fig. 122; Chen,
1989: 193, Figs. 32 c, d, Pl. VI 6.
Alox ornatum, Tan & Ng, 1995: 125, Fig. 10, pl. 5; Tan, 1996: 1022.
(see Tan & Ng, 1995: 125 for synonymy)
Material examined. – 1 female (CL 5.4 mm) (ZRC 2007.0530),
station T14, Maribohoc Bay, Bohol, 09°41.5'N 123°49.3'E, 101–110 m, mud with shells, 17 Jun.2004.

Colour. – Carapace white with bright orange marbling,
postfrontal pits and para-intestinal grooves dark orange
colour; chelipeds white, fingers bright red tipped with white,
pereiopodal carpi, propodi with orange spot proximally on
upper surface (Fig. 1C).
Distribution. – Australia, Indonesia, Moluccas, thePhilippines, Japan (Tan & Ng, 1995).

Arcania cornuta (MacGilchrist, 1905)(Fig. 1D)
Ixoides cornutus MacGilchrist, 1905: 255.
Ixoides cornutus, Serène & Vadon, 1981: 124; Chen, 1989: 227,
Pls. 1(11), 4(4), Fig. 21 a–c; Tan, 1996: 1033, Fig. 3 j–l, 4a–b.
Arcania cornuta, Galil, 2001a: 173, Figs. 1b, 4b; Komatsu et al.,2005: 106. (see Galil, 2001a: 173 for synonymy)
Material examined. – 1 male (CL 12.4 mm), 1 broken female (ZRC
2007.0531), station T28, Biking-Catamaran, Panglao, 09°35.0'N
123°51.4'E, 80 m, fine sand and mud, 1 Jul.2004.

Colour. – Carapace whitish, post-frontal pits and paraintestinal grooves darker; pereiopodal propodus with a brownspot proximally (Fig. 1B).

Distribution. – Fiji Islands, New Caledonia, Papua-New
Guinea, Japan, China, the Philippines, Vietnam, Persian Gulf,
Madagascar, Mozambique Channel (Galil, 2001a).

Remarks. – Alox chaunos, new species, is distinguished fromall congeners (see Tan & Ng, 1995) in having the male telson
distinctly longer than segment 6. All other Alox species, thetelson is subequal in length to segment 6 (see Tan & Ng,
1995; Naruse & Ng, 2006; present Fig. 5G). In generalcarapace form, A. chaunos is perhaps closer to A. rugosum(Stimpson, 1858) and A. uru Naruse & Ng, 2006. Alox
chaunos can easily be separated from A. uru by its lessprotruded front, lower gastric regions and more produced
hepatic region and a relatively longer and more slender G1
in which the median part is not visibly dilated and the tip
more tapering (Naruse & Ng, 2006: Figs. 1a, 2a, c, d, g, h).
Alox chaunos can be separated from A. rugosum by its moreeroded lateral carapace regions, more strongly produced
posterior margin of carapace and intestinal region, relatively
shorter and more elongate cheliped fingers, relatively stouterand shorter G1 (see Tan & Ng, 1995: Pls. 6D–F, 7, Fig. 12C,
H, I).

Arcania elongata Yokoya, 1933
(Fig. 1E)
Arcania undecimspinosa var. elongata Yokoya, 1933: 133, Fig. 47.
Arcania elongata, Tan, 1996: 1022, 1024, Figs. 1B, 2K–O. (seeGalil, 2001a: 176 for synonymy)
Material examined. – 2 males (CL 21.6 mm, 21.8 mm) (ZRC
2007.0532), Balicasag Island, tangle nets, Feb.2004; 3 males (ZRC
2007.0533), 2 Jun.2002. – 1 female (CL 11.7 mm) (MNHN), station
T19, Maribohoc Bay, Bohol, 09°42.2'N 123°50.8'E, 10–26m, mud,
20 Jun.2004.

Colour. – Carapace orange-red, slightly paler laterally,
posterior spines white; chelipeds orange-red proximally, paler
distally; pereiopodal meri pale orange, carpi, propodi, dactyli
white (Fig. 1E).

Etymology. – The species name is the Greek word for“porous, spongy”, chaunos, used as a noun in apposition.

Distribution. – New Caledonia, Australia, Japan, thePhilippines, South China Sea (Galil, 2001a).

Distribution. – Known only from type locations at Panglao
Island.

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Galil & Ng: Some Leucosiidae from the Philippines
Arcania gracilis (Henderson, 1893)(Fig. 1F)

Arcania gracilis, Galil 2001a: 184, Figs. 2c, 5d; Komatsu et al.,2005: 106. (see Galil 2001a: 184 for synonymy)

Arcania septemspinosa var. gracilis Henderson, 1893 : 403.
Arcania quinquespinosa, Serène & Vadon, 1981: 124; Chen, 1989:
208, Pl. I 7; Tan, 1996: 1022, 1028, Figs. 2b–e, 7f.

Material examined. – 1 male (CL 8.0 mm) (MNHN), station T6,
west of Baclayon, Bohol, 09°35.1'N 123°51.2'E, 34–82 m, sandymud with large sponges, 2 Jun.2004; 1 male (CL 10.4 mm) (ZRC

Fig. 1. Leucosiids from Panglao Island, the Philippines: A, Alox bothros, new species, holotype male (CL 9.0 mm) (NMCR); B, Alox chaunos,
new species, paratype female (CL 9.8 mm) (ZRC); C, Alox ornatum (Ihle, 1918), female (CL 5.4 mm) (ZRC 2007.0530); D, Arcania cornuta(MacGilchrist, 1905), male (CL 12.4 mm) (ZRC 2007.0531); E, Arcania elongata Yokoya, 1933, female (CL 11.7 mm) (MNHN); F, Arcania
gracilis (Henderson, 1893), male (CL 8.0 mm) (MNHN).

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THE RAFFLES BULLETIN OF ZOOLOGY 20072007.0534), station T26, Cortes, Bohol, 09°43.3'N 123°48.8'E, 123–
135 m, muddy, 24 Jun.2004; 1 female (CL 9.5 mm) (ZRC
2007.0535), station T33, Baclayon, Bohol, 09°36.0'N 123°53.7'E,
67–74 m, sand, 3 Jul.2004; 1 male (ZRC 2007.0536), station T23,
Cortes, Bohol, 09°42.2'N 123°50.6'E, 35–45 m, 21 Jun.2004; 1 male,
1 female (NMCR), station T28, Biking–Catarman, Panglao,
09°35.0'N 123°51.4'E, 80 m, muddy sand, 1 Jul.2004.

m, Jun.2002; 4 males (CL 18.2–18.3 mm), 1 female (ZRC
2007.0539), Mar.2004; 7 males, 3 ovigerous females, 7 females(ZRC 2007.0540), 20 Nov.2003; 4 males (ZRC 2007.0541), 20
Dec.2003; 1 male, 2 females (ZRC 2007.0542), May 2004; 1 male,
2 females (NMCR), May 2004; 6 males (CL 17.6–21.2 mm), 1
female (CL 21.1 mm), 2 ovigerous females (CL 21.5, 21.8 mm)(MNHN), Nov.2003; 2 males (CL 19.6, 20.4 mm), 1 ovigerous
female (CL 24.4 mm) (NMCR), Jan.2004; 1 female (CL 24.1 mm)(NMCR), Feb.2004; 1 female (CL 21.6 mm) (NMCR), Apr.2004;
2 males (ZRC 2007.0543), Apr. 2004; 1 male (CL 19.8 mm), 1
female (CL16.8 mm) (ZRC 2007.0544), 29 May.2004; 7 males (CL
18.5–20.5 mm 4 females (ZRC 2007.0545), station P3 (= P4),
09°31.1'N 123°41.5'E, ca. 100 m, 31 May 2004–2 Jul.2004; 1 male
(CL 19.9 mm) (MNHN), station P1, Maribohoc Bay, Bohol,
009°36.1'N 123°45.0'E, 150–200 m, 30 May.2004; 1 male (CL 20.1mm) (MNHN), station P2, Maribohoc Bay, Bohol, 09°39.0'N
123°43.8'E, 400 m, 30 May 2004; 6 males (CL 18.4–23.1 mm), 3
ovigerous females (CL 21.8–26.9 mm), 3 females (CL 21.1–25.5mm) (ZRC 2007.0547), 100–300 m, Maribohoc Bay, Bohol,
Nov.2003–Apr.2004; 2 males (CL 23.5 mm) (ZRC 2007.0546),
station P5, Pamilacan Island, 09°30.0'N 123°54.6'E, ca. 100 m, 3
Jun.2004; 1 male (ZRC 2007.0548), station T17, Cortes, Bohol,
09°41.8'N 123°49.1'E, muddy bottom with sponges, 132–137 m,
19 Jun.2004.

Color (young specimen). – Carapace orange-red, dotted withdark red spots anteriorly; cardiac, intestinal regions white,
red bar on cardiac region; cheliped meri rimmed with orangered, carpi, propodi paler distally; pereiopods pale. The “largebright red milk-white-edged ocellus” (Alcock, 1896: 266) socharacteristic of the adult, appears as a rectangular bar in theyoung (Fig. 1F).
Distribution. – Vanuatu, New Caledonia, Australia,
Indonesia, the Philippines, Japan, China, Singapore, India,
Sri Lanka, Laccadives, Persian Gulf, Madagascar, Red Sea(Galil, 2001a).

Arcania septemspinosa (Fabricius, 1787)(Fig. 2A)

Distribution. – Marquesas Islands, Loyalty Islands, Australia,
Japan, Korea, China, Taiwan, the Philippines, India,
Andamans, Seychelles, Mascarene Basin, South Africa (Galil,
2001a).

Cancer septemspinosus Fabricius, 1787: 325. (see Galil 2001a: 193
for synonymy)
Arcania heptacantha, Serène & Vadon, 1981: 124.
Arcania septemspinosa, Chen, 1989: 206, Fig. 9, Pl. II 6; Tan, 1996:
1022, 1029, Figs. 2a, 1d; Galil 2001a: 193, Figs. 3b, 7b. (seeGalil 2001a: 193 for synonymy)

Cateios frontalis (Miers, 1884)(Fig. 2B)

Material examined. – 1 male (CL 19.8 mm) (ZRC 2007.0537), 1
male (CL 15.2 mm) (NMCR), station T22, Cortes, Bohol, 09°42.5'N
123°50.7'E, 11-20 m, very muddy, 21 Jun.2004.

Oreophorus frontalis Miers, 1884: 254, Pl. 26 fig. B.
Oreophorus (Oreotlos) angulatus, Chen, 1989: 194, pl. VI 2.
Cateios frontalis, Tan & Ng, 1995: 135, Fig. 14, pl. 9b-f; Tan, 1996:
1022. (see Tan & Ng, 1995: 135 for synonymy)

Colour. – Carapace pinkish, with median carina and lateralspines orange-red; cheliped meri orange-red, carpi, propodi
pale orange; pereiopodal meri pale orange, carpi, propodi,
whitish, dactyli pale pink (Fig. 2A).

Material examined. – 1 male (CL 4.8 mm) (ZRC 2007.0549), 1
female (CL 5.6 mm) (MNHN), station T1, Bolod, Panglao,
09°32.4'N 123°47.3'E, 83–102 m, 30 May 2004.

Distribution. – Fiji, Australia, Indonesia, the Philippines, Gulfof Thailand, India, Madagascar, Mozambique Channel, South
Africa, Persian Gulf, Gulf of Aden, Red Sea (Galil, 2001a).

Color. – Carapace whitish, pale-orange laterally; cheliped
carpus with orange-red spot proximally on upper margin;
fingers with pale orange marbling (Fig. 2B).

Arcania undecimspinosa De Haan, 1841

Distribution. – Australia, Japan, Indonesia, the Philippines(Tan & Ng, 1995).

Arcania undecimspinosa De Haan, 1841: 135, Pl. 33 fig. 8.
Arcania undecimspinosa, Ihle, 1918: 265; Estampador, 1937: 512,
1959: 64; Serène & Vadon, 1981: 124; Chen, 1989: 204, fig 8,
Pl. II 4; Tan, 1996: 1022; Galil, 2001a: 197, Figs. 3D, 7D;
Komatsu et al., 2005: 106. (see Galil, 2001a: 197 for synonymy)

Euclosia crosnieri (Chen, 1989)(Fig. 2C)
Leucosia obtusifrons, Serène & Vadon, 1981: 125.
Leucosia crosnieri Chen, 1989: 236, Figs. 25, 26 a–e, Pl. I 8.
Euclosia crosnieri, Galil 2003c: 335, Figs. 1B, 3 B, 6B; Komatsu
et al., 2005: 106.

Material examined. – Balicasag Island, in deep slopes, with tangle
nets: 2 males, 1 ovigerous female (ZRC 2001.0397), Dec.2000; 4
males (CL 18.6–20.8 mm), 1 broken female (ZRC 2001.0398),
Dec.2000; 7 males (CL 15.4–21.4 mm), 6 females (CL 17.4–21.9mm) (ZRC 2001. 0576), 50–500 m, 28 Nov.2001; 10 males (CL
15.6–21.8 mm), 5 females (CL 19.2–25.2 mm), 3 ovigerous females(CL 20.4–23.2 mm) (ZRC 2001.0583), 50–500 m, 28 Nov.2001; 3
males (CL 21.4–23.3 mm), 5 females (CL 23.9–28.9 mm), 6
ovigerous females (CL 23.5–27.3 mm) (ZRC 2007.0538), 200–300

Material examined. – Maribohoc Bay, Bohol: 3 males, 1 female(ZRC 2007.0550), 200–300 m, tangle nets, Nov.2003–Mar.2004; 1
female (CL 21.9 mm) (ZRC 2007.0554), station P1, 009°36.1'N
123°45.0'E, 90–200 m, 30 May.2004. – Balicasag Island, tangle nets:
1 female (CL 30.2 mm) (ZRC 2007.0555), station P3 (= P4),

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Galil & Ng: Some Leucosiidae from the Philippines
09°31.1'N 123°41.5'E, 100 m, 31 May.2004; 1 female (ZRC
2007.0556), Jun.2002; 17 males (CL 21.8–32.3 mm), 4 ovigerous
females (CL 29.6–32.1 mm), 14 females (CL 20.3–31.6 mm) (ZRC
2001.0565), 50–500 m, 28 Nov.2001; 9 males, 14 females (ZRC
2007.0551), Jun.2002; 3 males (CL 30.6, 31.1, 30.0 mm), 2 females(CL 29.7, 29.9 mm) (ZRC 2007.0552), Nov.2003; 3 males (CL 30.632.3 mm), 1 female (CL 29.9 mm) (MNHN), 100–300 m, 11 Mar.–
4 Apr.2004; 2 males (CL 22.8, 32.1 mm) (NMCR), 2 Mar.2004; 1
male (CL 31.0 mm), 1 ovigerous female (CL 31.2 mm), 1 juvenile
female (CL 21.3 mm) (NMCR), May.2004; 1 ovigerous female (CL32.6 mm) (MNHN), 28 May.2004; 1 male (CL 31.6 mm), 1 female(CL 30.2 mm) (ZRC 2007.0553), 29 May.2004.

Iphiculus spongiosus Adams & White, 1848
Iphiculus spongiosus Adams & White, 1848: 57, Pl. 15 Fig. 5;
Estampador, 1937: 512; Estampador, 1959: 64; Serène & Vadon,
1981: 124; Chen, 1989: 233, Figs. 4 c–f, Pl. IV 5. (see Chen &Sun, 2002: 372 for synonymy)
Iphiclus spongiosus, Tan, 1996: 1022. (incorrect spelling)
Material examined. – 3 males (CL 10.4–11.2 mm), 2 females (CL10.7 mm) (ZRC 2007.0563), station T10, off San Isidro, Panglao,
09°33.4'/09°33.8'N 123°49.6'/123°51.5'E, 117–124 m, fine sand, 15
Jun.2004; 3 males (CL 6.3–7.4 mm), 1 female (CL 11.3 mm)(MNHN), station T19, Cortes, Bohol, 09°42.2'N 123°50.8'E, 10–
26 m, muddy bottom, 20 Jun.2004; 3 males (CL 8.6–12.0 mm) (ZRC
2007.0564), station T22, Cortes, Bohol, 09°42.5'N 123°50.7'E, 11–
20 m, muddy bottom, 21 Jun.2004; 1 male (ZRC 2007.0565), station
T25, Cortes, Bohol, 09°41.1'N 123°49.3'E, 160–210 m, fine sandand mud, 24 Jun.2004; 1 male (CL10.1 mm), 1 female (CL 10.4mm) (NMCR), station T27, between Panglao and Pamilacan Islands,
09°33.4'N 123°51.0'E, 106–137 m, fine sand and mud, 21 Jun.2004;
1 male (ZRC 2007.0566), station T7, west of Baclayon, Bohol,
09°36.1'N 123°53.3'E, muddy fine sand, 61–62m, 3 Jun.2004; 1
male, 1 female (NMCR), station T24, 09°42.3'N 123°50.3'E, mud
and wood, 35–57 m, 23 Jun.2004.

Colour. – “In young specimens …. there are many obliqueand longitudinal stripes which are not present in full grownspecimens” (Chen, 1989: 236). Carapace of young specimen
pinkish, with deep orange lines; cheliped merus, carpus andproximal part palm and fingers reddish; pereiopods whiteringed in bright red (Fig. 2C).
Distribution. – Australia, Indonesia, the Philippines (Galil,
2003c).

Distribution. – Arafura Sea, Indonesia, the Philippines, China
Seas, Japan, Gulf of Thailand, Singapore, Bay of Bengal, Red
Sea (Chen, 1989).

Euclosia scitula Galil, 2003
Euclosia scitula Galil, 2003c: 339, Figs. 2a, 4c, 6g; Komatsu et al.,2005: 106.
Material examined. – Balicasag Island, tangle nets: 2 males (CL27.2, 27.0 mm), 3 juvenile females (CL 20.5–20.9 mm) (ZRC
2001.0574), 50–500 m, 28 Nov.2001; 4 males (CL 26.5–28.0 mm),
4 females (CL 26.4–27.7 mm) (NMCR), 200–300 m, Jun.2002; 2
males (CL 21.0, 30.1 mm) (MNHN), Nov.2003; 4 males, 1 female(ZRC 2007.0558), Jan.–May 2004; 1 male, 1 female (ZRC
2007.0559), Mar.2004; 3 males (CL 26.1–28.9 mm), 1 female (CL24.4 mm) (MNHN), 2 Mar.2004; 3 males (CL 28.5–30.2 mm)(MNHN), Apr.2004; 1 male (CL 31.3 mm), 1 female (CL 30.3 mm),
2 juveniles (CL 20.2, 20.9 mm) (MNHN), 29 May.2004; 2 males(CL 29.3, 30.7 mm) (ZRC 2007.0560), station P3 (= P4), 09°31.1'N
123°41.5'E, 100 m, 31 May.2004; 1 female (ZRC 2007.0561), station
P3 (= P4), 09°31.1'N 123°41.5'E, 100 m, 31 May.2004; 2 males(ZRC 2007.0557), 100–300 m, Maribohoc Bay, Bohol, tangle nets,
Nov.2003–Apr.2004.

Leucosia craniolaris (Linnaeus, 1758)(Fig. 2D)
Cancer craniolaris Linnaeus, 1758: 626.
Leucosia perlata, Estampador, 1937: 512; Estampador, 1959: 62;
Tan, 1996: 1023, 1037, Figs. 5b–e.
Leucosia craniolaris, Galil, 2003b: 184, Figs. 1a, 2a, b. (see Galil,
2003b: 184 for syonymy)
Material examined. – 1 male (CL 19.7 mm) (MNHN), station M11,
Sungcolan Bay, Panglao, 09°38.3'N 123°49.6'E, 0–3 m, rocks,
mangrove, seagrasses, 6 Jun.2004; 1 male (CL 19.0 mm) (ZRC
2007.0567), station M3 (= M25), Danao, Panglao, 09°32.5'N/
09°33.1'N 123°44.7'E/123°45.5'E, intertidal, muddy-sand,seagrasses, 31 May – 17 Jun.2004.

Distribution. – The Philippines, Thailand (Galil, 2003c).

Colour. – Carapace whitish, marbled anteriorly with palekhaki color, two pairs of pale spots laterally on gastric regions,a pair of prominent rust-coloured spots near base of last pairof legs. Chelipeds and pereiopods pale, pale rust spots distallyon pereiopodal meri (Fig. 2D).

Euclosia unidentata (De Haan, 1841)
Leucosia unidentata De Haan, 1841: 133, Pl., 33 Fig. 3.
Euclosia unidentata, Komatsu et al., 2005: 106. (see Galil, 2003c:
342 for synonymy)

Distribution. – Palau Islands, Caroline Islands, Ryukyu
Islands, Indonesia, the Philippines, Malaysia (Galil, 2003b).

Material examined. – Balicasag Island, tangle nets: 2 females (CL28.9, 20.5 mm) (ZRC 2001.0574), 50–500 m, 28 Nov.2001; 1 male
(CL 28.8 mm) (ZRC 2007.0562), 200–300 m, Jun.2002; 1 male
(CL 31.6 mm), 1 ovigerous female (CL 29.5 mm) (NMCR),
Nov.2003.

Leucosia punctata Bell, 1855
Leucosia punctata Bell, 1855a: 362; Estampador, 1937: 512;
Estampador, 1959: 62; Galil, 2003b: 187, Figs. 1c, 2e, f. (seeGalil, 2003b: 187 for synonymy)

Distribution. – Japan, China, Gulf of Tonkin, Taiwan,
Indonesia (Galil, 2003c), the Philippines (Komatsu et al.,2005).

Material examined. – 1 male (CL 19.0 mm) (NMCR), station D12,
channel between Tagbilaran town (Bohol) and Panglao, 09°38.5'N

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123°51.0'E, 2–4m, muddy bottom, shells and asteroids, 28 Jun.2004.
– 2 males (CL 19.5, 14.1 mm), 1 female (CL 16.4 mm) (ZRC
2007.0568), station D13, Tagbilaran channel, 09°38.0'N 123°51.4'E,
2–3m, sandy bottom, oyster shells, 29 Jun.2004.

m, dead coral, 21 Jun.2004; 1 male (NMCR), station T23, Cortes,
Panglao, 09°42.2'N 123°50.6'E, 35–45 m, black mud, 21 Jun.2004.

Colour. – Carapace ruby-red but for tip of posterior spine;cheliped whitish but for red merus; pereiopods reddish
proximally, pale distally (Fig. 2F).

Colour. – “Greyish-brown, with two darker spots on the
branchial regions” (Bell, 1855b: 287).
Distribution. – Australia, Indonesia, the Philippines (Galil,
2003b).

Distribution. – Australia, Papua New Guinea, Indonesia, thePhilippines, South China Sea, Gulf of Thailand, Myanmar,
Bay of Bengal (Galil, 2001b).

Leucosia rubripalma Galil, 2003
(Fig. 2E)

Myra eudactyla (Bell, 1855)
Myrodes eudactylus Bell, 1855a: 364; Bell, 1855b: 299, Pl. 32 Fig.
6; Bell, 1855c: 13; Estampador, 1937: 512; Estampador, 1959:
63; Tan, 1996: 1023, 1044.
Myra eudactyla, Galil, 2001b: 425, Figs. 2b, 11; Komatsu et al.,2005: 106.

Leucosia rubripalma Galil, 2003b: 188, Figs. 1d, 2g, h.
Material examined. – 1 male (CL 21.2 mm) (ZRC 2007.0569),
station R24, Bingag, Panglao, coral platform, 09°37.5'N 123°46.8'E,
0–2 m, coral rocks, 6 Jun.2004.

Material examined. – Balicasag Island, tangle nets: 2 males (CL
32.1–30.7 mm), 1 female (CL 38.5 mm), 1 ovigerous female (CL41.4 mm) (ZRC 2007.0572), 200–300 m, Jun.2002; 1 male (CL39.0 mm) (ZRC 2001.0572), 50–500 m, 28 Nov.2001. Singapore:
1 male (ZRC 1998.1217), off Katong, Singapore, dredge, coll. P.
K. L. Ng, 1990s.

Colour. – Carapace ochre-coloured, paler posteriorly, a pairof red spots on the branchial regions, a pair of prominentorange-coloured spots near base of the last pair of legs.
Chelipeds pale ochre, with bright-orange patch on innersurface of palm. Pereiopods white, orange spots distally on
upper margin of meri 1-3, proximally on propodi 1–3 (Fig.
2E).

Colour. – The Singapore specimen (ZRC 1998.1217) wasan uneven chocolate-brown on all dorsal surfaces when fresh,with the ventral surfaces dirty white.

Distribution. – New Caledonia, Indonesia, Singapore (Galil,
2003b), the Philippines (new record).

Distribution. – New Caledonia, Australia, Torres Straits, New
Guinea, Indonesia, the Philippines, Tonkin Bay, Gulf ofThailand, Andamans, Gulf of Aden (Galil, 2001b). A
specimen (ZRC 1998.1217) collected represents a new record
for Singapore.

Myra curtimana Galil, 2001
Myra curtimana Galil, 2001b: 421, Figs. 1f, 9; Komatsu et al., 2005:
106.
Material examined. – Balicasag Island, tangle nets: 1 female (CL24.9 mm) (ZRC 2001.0405), Dec.2000; 1 male (CL 25.8 mm), 1
female (CL 27.1 mm) (ZRC 2001.0407), Dec.2000; 2 males (CL21.2, 19.5 mm), 3 females (CL 30.1–32.2 mm) (ZRC 2001.0569),
50–500 m, 28 Nov.2001; 2 females (CL 31.5, 26.8 mm) (ZRC
2001.0570), 50–500 m, 28 Nov.2001; 4 males (CL 23.6–25.3 mm)
1 female (NMCR), 200–300 m, Jun.2002; 1 male (MNHN),
May.2004.

Myra grandis Zarenkov, 1990
Myra grandis Zarenkov, 1990: 65, pl. 6, Figs. 8–12; Galil, 2001b:
429, Figs. 2d, 13; Komatsu et al., 2005: 106, 109, Fig. 1E, 2a–c.
Material examined. – Balicasag Island, tangle nets: 1 male (CL30.8 mm) (ZRC 2001.0403), Dec 2000; 1 male (CL 32.9 mm) (ZRC
2001.0561), 50–500 m, 28 Nov.2001; 1 male (CL 38.6 mm), 1
female (CL 42.1 mm) (ZRC 2001.0575), 50–500 m, 28 Nov. 2001;
1 female (CL 43.1 mm) (NMCR), 200–300 m, Jun. 2002.

Distribution. – Fiji, New Caledonia, Australia, Indonesia, thePhilippines (Galil, 2001b).

Distribution. – Marquesas Islands, Madagascar, Kenya, thePhilippines (Galil, 2001b; Komatsu et al., 2005).

Myra elegans Bell, 1855
(Fig. 2F)
Myra elegans Bell, 1855a: 364; Serène & Vadon, 1981: 124; Chen,
1989: 223, Figs. 19, Pl. I 12; Tan, 1996: 1023, 1043; Galil,
2001b: 423, Figs. 2a, 10. (see Galil, 2001b: 423 for synonymy)

Myra tumidospina Galil, 2001
(Fig. 3A)
Myra tumidospina Galil, 2001b: 433, Figs. 3b, 17.

Material examined. – 1 male (CL 15.2 mm), 4 ovigerous females(CL 16.2–18.2 mm) (ZRC 2007.0570), station T19, Cortes, Bohol,
09°42.2'N 123°50.8'E, 10–26 m, muddy, 20 Jun.2004; 1 male (CL15.8 mm) (ZRC 2007.0571), station T21, Cortes, Bohol, 09°42.8'N
123°50.6'E, 12 m, dead coral, 21 Jun.2004; 1 male (CL 18.0 mm),
(MNHN) station T22, Cortes, Bohol, 09°42.5'N 123°50.7'E, 11–20

Material examined. – Balicasag Island, tangle nets: 2 females (CL27.3, 31.6 mm) (ZRC 2001.0407), Dec.2000; 15 males (CL 20.9–
29.9 mm), 2 females (CL 28.4, 28.5 mm) (ZRC 2001.0570), 50–
500 m; 14 males (CL 21.0–29.9 mm), 9 females (CL 23.6–32.9 mm)

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Galil & Ng: Some Leucosiidae from the Philippines(MNHN), 200–300 m, Jun.2002; 2 females (CL 25.1, 27.3 mm), 2
males (CL 25.6, 27.0 mm) (NMCR), Mar.2004; 2 males (CL 29.1,
26.4 mm) (ZRC 2007.0573), 2 Mar.2004; 2 males (CL 23.1–24.4mm) (ZRC 2007.0574), 29 May.2004; 1 male (NMCR), Dec.2003;
1 male (MNHN), Maribohoc Bay, Bohol, Nov.2003–Apr.2004; 1

juvenile (CL 10.6 mm) (ZRC 2007.0575), station T6, west of
Baclayon, Bohol, 09°35.1'N 123°51.2'E, coarse muddy sand withlarge sponges, 34–82 m, 2 Jun.2004.

Colour. – See Fig. 3A.

Fig. 2. Leucosiids from Panglao Island, the Philippines: A, Arcania septemspinosa (Fabricius, 1787), male (CL 19.8 mm) (ZRC 2007.0537);
B, Cateios frontalis (Miers, 1884), male (CL 4.8 mm) (ZRC 2007.0549); C, Euclosia crosnieri (Chen, 1989), 1 (CL 21.9 mm) (ZRC
2007.0554); D, Leucosia craniolaris (Linnaeus, 1758), male (CL 19.7 mm) (MNHN); E, Leucosia rubripalma Galil, 2003, 1(CL 21.2mm) (ZRC 2007.0569); F, Myra elegans Bell, 1855, male (CL 15.2 mm) (ZRC 2007.0570).

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Pariphiculus coronatus (Alcock & Anderson, 1894)

Distribution. – Fiji, New Caledonia, Vanuatu, Indonesia(Galil, 2001b), the Philippines (new record).

Pariphiculus coronatus, Serène & Vadon, 1981: 124; Chen, 1989:
229, Fig. 22, Pls. I 3, III 5; Tan, 1996: 1023, 1048; Komatsu etal., 2005: 106.

Parilia major Sakai, 1961

Material examined. – Balicasag Island, tangle nets: 2 males (CL23.3, 24.3 mm), 4 females (CL 23.4–24.5 mm), 1 ovigerous female(CL 26.3 mm) (ZRC 2001.0571), 28 Nov.2001; 1 male (CL 24.3mm), 1 female (CL 26.5 mm), ZRC 2001.0573, 28 Nov.2001; 1
female (CL 25.8 mm) (ZRC 2001.0339), Dec.2000, 3 males (CL
23.7–25.6 mm), 4 females (CL 19.7–25.9 mm) (ZRC 2007.0591),
Jun.2002; 1 male (CL 24.9 mm), 1 female (CL 25.3 mm) (ZRC
2007.0592), 2 Mar.2004; 1 male, 1 female (NMCR), Nov.2004, 2
males, 1 female (MNHN), Nov.2004; 3 males, 2 females, 1 ovigerous
female (ZRC 2007.0593), Feb.2004; 1 male, 1 female (ZRC
2007.0594), Mar.2004; 1 juvenile (CL 6.9 mm) (ZRC 2007.0595),
T25, Cortes, Bohol, 09°41.1'N 123°49.3'E, fine sand and mud, 160–
210 m, 24 Jun.2004.

Parilia major Sakai, 1961: 137, Pl. 3; Serène & Vadon, 1981:124;
Chen, 1989: 233, Fig. 24, Pl. II 1; Tan, 1996: 1023, 1046, Figs.
7 f–k; Komatsu et al., 2005: 106. (see Chen & Sun, 2002: 365,
for synonymy)
Material examined. – Balicasag Island, tangle nets: 1 male (CL43.7 mm), 1 ovigerous female (CL 46.7 mm), 1 female (CL 46.3mm) (ZRC 2001.0406), Dec. 2000; 2 males, 2 ovigerous females,
4 females (ZRC 2001.0370), Dec.2000; 3 females (CL 21.9–30.6mm) (ZRC 2007.0576), Dec.2003; 3 males (CL 30.3-46.9 mm), 3
females (CL 43.8–45.8 mm) (ZRC 2007.0577), Nov.2003; 4 females(CL 31.1–44.7 mm) (ZRC 2007.0580), Mar.2004; 1 ovigerous
female (CL 42.3 mm) (ZRC 2007.0581), May.2004; 4 males (CL
44.7–61.3 mm), 4 ovigerous females (CL 42.4–46.8 mm), 4 females(CL 30.4–44.2 mm) (ZRC 2007.0582), 28 May.2004; 2 males, 2
females (NMCR), Balicasag Island, Feb.2004; 1 male, 2 females(ZRC 2007.0585), Jan.2004; 4 males (CL 47.8–65.8 mm), 2 females(CL 41.6, 43.6 mm) (ZRC 2007.0583), 300 m, soft bottom withsponges, 27 May.2004; 3 males, 3 females (ZRC 2007.0578), station
P2, Maribohoc Bay, Bohol, 09°39.0'N 123°43.8'E, Oct.2003; 32
males, 22 ovigerous females, 19 females. (ZRC 2007.0579),
Maribohoc Bay, Bohol, 100–300 m, tangle nets, Nov.2003–
Apr.2004; 2 males (CL 63.9, 58.9 mm) (ZRC 2007.0584), Momo
Beach, Panglao, 350 m, tangle net, 2 Jun.2004; 5 males (CL 42.5–
61.4 mm), 1 ovigerous female (CL 43.0 mm) (MNHN), 7 Jul.2004;
1 female (ZRC 2007.0586), station P1, Maribohoc Bay, Bohol,
009°36.1'N 123°45.0'E, 90–200 m, 30 May2004; 2 males, 1 female(ZRC 2007.0587), station P1, Maribohoc Bay, Bohol, 009°36.1'N
123°45.0'E, 90–200 m, 30 May.2004; 4 males (MNHN), station P2,
Maribohoc Bay, Bohol, 009°36.1'N 123°45.0'E, 400 m, 30
May.2004.

Distribution. – Indonesia, the Philippines, China Seas,
Vietnam, Japan, Bay of Bengal, Persian Gulf, Red Sea (Tan,
1996).

Pariphiculus mariannae (Herklots, 1852)
Ilia mariannae Herklots, 1852: 36, Fig. 2.
Pariphiculus mariannae, Serène & Vadon, 1981: 124; Chen, 1989:
231, Figs. 23, Pl. IV 3; Tan, 1996: 1023, 1049.
Material examined. – 1 female (CL 30.1 mm) (ZRC 2007.0596),
Balicasag Island, 84-95 m, tangle nets, 26 Oct. 2003.

Distribution. – Indonesia, the Philippines, South China Sea,
Myanmar, India, Arabian Sea (Tan, 1996, Tirmizi & Kazmi,1988).

Distribution. – Indonesia, the Philippines, Japan (Tan, 1996).
Praebebalia septemspinosa Sakai, 1983
Pariphiculus agariciferus Ihle, 1918
(Fig. 3B)

Praebebalia septemspinosa Sakai, 1983: 625, Figs. 2d–h; Chen,
1989: 192; Tan, 1996: 1023, 1050, Figs. 6k–m; Galil, 2001d:
273, Figs. 4a–c, 5d; Komatsu et al., 2005: 106.

Pariphiculus agariciferus Ihle, 1918: 250, Fig. 230; Serène & Vadon,
1981: 124; Chen, 1989: 231, Figs. 21 d-g, Pl. II 5; Tan, 1996:
1023; Komatsu et al., 2005: 106. (see Chen & Sun, 2002: 374,
for synonymy)

Material examined. – Balicasag Island, tangle nets: 3 males (CL
13.5–13.8 mm), 1 ovigerous female (CL 13.5 mm) (ZRC
2001.0402), Dec. 2000; 4 males (CL 13.2–14.8 mm) (ZRC
2001.0557), 50–500 m, 28 Nov.2001; 1 ovigerous female (CL 12.5mm) (ZRC 2001.0558), 50–500 m, 28 Nov.2001; 2 females (CL12.7, 14 mm) (NMCR), 200–300 m, Jun.2002; 1 male (CL 13.8mm), 1 ovigerous female (CL 14.2 mm) (ZRC 2007.0598), 2
Mar.2004; 1 male (MNHN), Nov.2003; 15 males (CL 11.5–15.5mm), 5 ovigerous females (CL 12.5 mm) (ZRC 2001.0558), 50–
500 m, 28 Nov.2001; 2 females (CL 11.5–13.5 mm) (ZRC
2007.0597), Maribohoc Bay, Bohol, 100–300 m, Nov.2003–
Apr.2004; 2 males, 1 ovigerous female (ZRC 2007.0599), station
P1, 009°36.1'N 123°45.0'E, 30 May.2004; 3 males, 1 ovigerous
female (MNHN), Maribohoc Bay, Bohol, Nov.2003.

Material examined. – Balicasag Island, tangle nets: 1 female (CL18.1mm) (MNHN), station P2, 9º0.039'N 123º0.044'E, 400 m, 30
May.2004; 1 female (NMCR), station P1, 30 May.2004; 1 male
(MNHN), station P3 (= P4), 2 Jul.2004; 3 males, 1 female (ZRC
2007.0588), Nov.2003; 1 male (ZRC 2007.0589), May.2004; 1 male,
1 female (MNHN), Jun.2002; 1 male, 1 female (ZRC 2001.0559),
28 Nov.2001; 1 male, 1 female (ZRC 2007.0590), 2 Mar.2004.

Colour. – Carapace and chelipeds dull red, granules somewhat
paler; pereiopods dull red but for white ring distally on merus(Fig. 3B).

Distribution. – The Philippines (Sakai, 1983; Komatsu et al.,2005).

Distribution. – Indonesia, the Philippines, South China Sea,
Japan (Chen, 1989).

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Galil & Ng: Some Leucosiidae from the Philippines
Remarks. – This species was described on the basis of one
male and one female from Balicasag Island, and we add
another female to the known specimen list. The species,
according to Komatsu et al. (2005) is intermediate between
Raylilia mirabilis (Zarenkov, 1990) and R. coniculifera Galil,
2001c.

Raylilia intermedia Komatsu, Manuel & Takeda, 2005
Raylilia intermedia Komatsu, Manuel & Takeda, 2005: 109, Figs.
3, 4, 8A, B.
Material examined. – 1 female (CL 13.1 mm) (ZRC 2007.0600),
Balicasag Island, tangle nets, Feb. 2004.

Fig. 3. Leucosiids from Panglao Island, the Philippines: A, Myra tumidospina Galil, 2001, juvenile (CL 10.6 mm) (ZRC 2007.0575); B,
Pariphiculus agariciferus Ihle, 1918, female (CL 18.1mm) (MNHN); C, Urnalana cumingii (Bell, 1855), male (CL 9.4 mm) (ZRC 2007.0624);
D, Urnalana cristata, new species, holotype male (CL 11.1 mm) (NMCR); E, Urnalana pulchella (Bell, 1855), male (CL 9.2 mm) (ZRC
2007.0628); F, Urnalana whitei (Bell, 1855), male (CL 8.0 mm) (ZRC 2007.0629).

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Material examined. – Balicasag Island, tangle nets: 2 males (ZRC
2001.0564), 28 Nov.2001; 19 males (CL 16.6–28.1 mm), 14
ovigerous females (CL 16.0–20.5 mm) (ZRC 2001.0566), 50–500m, 28 Nov.2001; 1 male, 4 females (ZRC 2001.0371), Dec.2000;
3 males, 1 ovigerous female (MNHN), Nov.2003; 4 males (CL 18.1–
19.2 mm) (ZRC 2007.0608), Dec.2003; 5 males (CL 17.2–19.3 mm),
1 ovigerous female (CL 17.9 mm) (ZRC 2007.0609), station P2, 20
Jan.2004; 11 males (CL 15.2–20.6 mm), 2 ovigerous females (CL20.7 mm) (NMCR), Mar.2004; 1 male (ZRC 2007.0610), 2
Mar.2004; 3 males, 3 ovigerous females(ZRC 2007.0611),
May.2004; 1 male, 3 ovigerous females (MNHN), Feb.2004; 1 male
(ZRC 2007.0614), Jan.2004; 1 ovigerous female (ZRC 2007.0615),
28 May.2004; 1 male, 3 ovigerous females (ZRC 2007.0616), station
P4, 9º0.031'N 123º0.041'E, 100 m; 4 males, 1 ovigerous female(ZRC 2007.0617), station P3 (= P4), 09°31.1'N 123°41.5'E, 31
May.2004; 1 male (ZRC 2007.0618), station P3 (= P4), 09°31.1'N
123°41.5'E, 31 May.2004; 2 males, 1 female (ZRC 2007.0619),
station P3 (=P4), 09°31.1'N 123°41.5'E, 31 May.2004; 1 male (ZRC
2007.0612), station P1, Maribohoc Bay, Bohol, 09°36.1'N
123°45.0'E, 150–200 m, 30 May.2004; 1 male, 1 ovigerous female(ZRC 2007.0613), Maribohoc Bay, Bohol, Nov.2003–Apr.2004; 1
ovigerous female, 1 female (ZRC 2007.0620), station P5, Pamilacan
Island, 09°30.0'N 123°54.6'E, tangle nets from local fishermen, ca.
100 m, 3 Jun.2004; 2 females (ZRC 2007.0621), station T10, off
San Isidro, Panglao, 09°33.4'N/09°33.8'N 123°49.6'E/123°51.5'E,
117–124 m, fine sand, 15 Jun.2004; 5 males, 1 juvenile (ZRC
2007.0622), station T27, between Panglao and Pamilacan islands,
09°33.4'N 123°51.0'E, 106–137 m, fine sand and mud, 21 Jun.2004.
Comparative material. Tokoyo eburnea: 13 males (CL 14.7–19.1mm), 9 ovigerous females (CL 14.8–18.7 mm) (ZRC 2001.0567),
Tomioka, Amakusa, Kyushu, Japan, coll. J. C. Y. Lai & S. Arakaki,
3 Sep.2002.

Distribution. – Known only from the Philippines thus far(Komatsu et al., 2005).

Seulocia cristata Galil, 2005
Leucosia rhomboidalis, Chen, 1989: 240, Fig. 30 b-c.
Seulocia cristata Galil, 2005b: 46, Fig. 1c.
Material examined. – 2 females (CL 13.9–14.1 mm) (NMCR),
station R52, Bingag, Panglao, 09°37.4'N 123°46.9'E, 3–20 m, coral
plateau with sand, 19 Jun.2004; 5 males (CL 9.2–13.3 mm), 4
females (CL 13.1–13.4 mm) (ZRC 2007.0601), station T19, Cortes,
Bohol, 09°42.2'N 123°50.8'E, 10–26 m, muddy, 20 Jun.2004; 2
males (CL 12.8–13.7 mm) (MNHN), station T21, Cortes, Bohol,
09°42.8'N 123°50.6'E, 12 m, dead coral, 21 Jun.2004; 2 males (CL12.9, 13.4 mm), 2 ovigerous females (CL 13.5, 13.7 mm), 1 female(CL 12.6 mm) (ZRC 2007.0602), station T22, Cortes, Bohol,
09°42.5'N 123°50.7'E, 11–20 m, dead coral, 21 Jun.2004; 2 males(CL 12.2, 12.3 mm), 1 ovigerous female (CL 13.0 mm), 1 female(CL 13.3 mm) (ZRC 2007.0603), station T23, Cortes, Bohol,
09°42.2'N 123°50.6'E, 35–45 m, black mud, 21 Jun.2004; 1 female(MNHN), station S20, Manga, Bohol, 09°41.8'N 123°51.1'E, mud,
8–10 m, 20 Jun.2004.

Distribution. – Indonesia, the Philippines, Japan, Singapore,
Thailand (Galil, 2005b).

Tanaoa pustulosus(Wood-Mason, in Wood-Mason & Alcock, 1891)

Remarks. – Komatsu et al. (2005: 116) described a new
species from the Philippines, T. trilobata, which they argued
is very close to T. eburnea but differing in three main
characters: the posterior lobes on the carapace margin are
rounded (vs. triangular in T. eburnea), the space between the

Randallia pustulosa Wood-Mason, in Wood-Mason & Alcock,
1891: 246;
Serène & Vadon, 1981: 124; Chen, 1989: 217, Fig. 15, Pl. IV 1;
Tan, 1996: 1023, 1054.
Tanaoa pustulosus, Galil, 2003a: 404, Figs. 1d, 3g, h; Komatsu etal., 2005: 106. (see Galil, 2003a: 404 for synonymy)
Material examined. – Balicasag Island, tangle nets: 3 males (CL
33.4–34.7 mm), 3 females (1 parasitized) (CL 28.3–34.8 mm), 1
ovigerous female (CL 34.5 mm) (ZRC 2001.0567), 50–500 m, 28
Nov.2001; 1 male, 1 ovigerous female (CL 29.7 mm) (ZRC
2007.0604), 100–300 m, Nov.2003–Apr.2004; 2 males (CL 30.8,
32.1 mm) (ZRC 2007.0605), Mar.2004; 1 female (CL 31.4 mm)(ZRC 2007.0606), 2 Mar.2004; 1 male (CL 33.7 mm) (ZRC
2007.0607), 28 May.2004.

Distribution. – Fiji, New Caledonia, New Zealand, Caroline
Islands, Japan, Taiwan, Indonesia, the Philippines, Andaman
Sea, Laccadive Sea, Seychelles, Comoro Islands, Madagascar,
East Africa (Galil, 2003a).

Tokoyo eburnea (Alcock, 1896)(Fig. 4)
Randallia eburnea Alcock, 1896: 197; Serène & Vadon, 1981: 124;
Chen, 1989: 212, Figs. 12, 13.
Tokoyo eburnea, Galil, 2003a: 408, Figs. 1f, 4d–f; Komatsu et al.,2005: 106, Figs. 7d–f, h, 8D. (see Galil, 2003a: 408 forsynonymy)
Tokoyo trilobata Komatsu, Manuel & Takeda, 2005: 116, Figs. 5,6, 7a–c, g, 8C, D.

Fig. 4. Posterior carapace margins of Tokoyo eburnea showing
variation. ZRC 2001.0567, Kyushu, Japan, coll. J. C. Y. Lai & S.
Arakaki, 3 Sep.2002. A, sharp teeth, male (CL 18.4 mm); rounded
teeth, male (CL 18.4 mm); fused teeth, male (CL 18.1 mm).

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Galil & Ng: Some Leucosiidae from the Philippines
lengthwise on endopod of female. Hepatic region bearing oval
tumescence, branchial region bearing elongate tumescences.
Lateral angle of carapace prominent, overhanging thoracic
sinus, margin smooth. Thoracic sinus deep, setose, anteriorly
defined by overhanging margin of pterygostomian region;row of granules ventrally. Epimeral ledge visible in dorsal
view, continuous with posterior margin, margin beaded.
Posterior margin prominent, beaded. Chelipeds subequal,robust. Cheliped merus half as long as carapace; entirely
granular but for smooth patch medially on lower surface;anterior, posterior margins bearing conic granules, larger
medially; upper surface bearing setose patch proximally.
Carpus with row of granules on inner margin. Upper marginof palm with prominent, smooth carina; lower inner margin
with scalloped carina extending to proximal part of pollex;
outer surface of palm proximally with row of minute granules,
parallel with lower margin. Upper margin of dactyl carinate.
Pereiopodal meri 1–3 with granulate rows on lower surface,
distally carinate on upper surface; upper and lower margins
of merus of fourth pereiopod prominently carinate.
Pereiopodal carpi prominently keeled dorsally; propodi keeled
dorsally, ventrally. Male abdominal sulcus deep, nearlyreaching buccal cavity; lateral margin bearing distinct ridge
fitting into suture between abdominal segments. Male
abdomen with segment 2 small; segments 3–5 fused,
proximally with median furrow; segment 6 large, trapezoid,
medially denticulate; telson triangular. Shaft of G1 short,stout, sinuous, distally setose; cornuted apical process curved,
sigmoid. G2 short, curved, apex scoop-like.

posterior carapace lobes was relatively wider (closer together
in T. eburnea) and the tip of the G1 was more prominently
bent upwards. Alcock (1896) described T. eburnea (asRandallia eburnea) from the Bay of Bengal. His illustrationof the species (in Alcock & Anderson, 1897: Pl. 30 Fig. 4)
clearly shows the rounded and well spaced lobes on theposterior margin that Komatsu et al. (2005) maintain is adistinguishing character of T. trilobata. Comparison of theillustrations provided by Komatsu et al. (2005: Figs. 5, 6,
7a–c, g, 8c, d) with a large series of specimens of T. eburnea
from Japan, Taiwan, Indonesia, Philippines and western
Thailand, convinced us that all the differences can easily be
explained by variation. The form of the posterior carapace
lobes varies considerably, from rounded to broadly triangular
to almost triangular, the tip varying from sharp to round (Fig.
4A, B). The distance between the lobes also varies, from far
to close, and in a few cases, they are almost completely fused,
forming one lobiform structure (Fig. 4C). With regards tothe structure of the tip of the G1, it is usually gently curved
upwards (proximally) but in a few specimens, the curvature
is more pronounced. The form of the G1 cannot be correlatedwith the other characters, and it is clear the tip varies morethan previously recognized. Komatsu kindly provided us
colour figures of what he regarded were two species with thefollowing comments: “We found two color types of “Tokoyo
eburnea” in the Balicasag collection. One is larger andspeckled (= T. eburnea), another is simply reddish (= T.
trilobata). Since there are small morphological differences
between them, we described one as new. But that may be arough and ready conclusion.” (H. Komatsu, in litt. to P. K.
L. Ng). Having examined a large series, including many live
as well as freshly preserved specimens from Balicasag andelsewhere, we believe that these differences are well within
the colour variation we have observed. In summary, we areof the opinion that T. trilobata is synonymous with T.
eburnea.

Colour. – See Fig. 3D.
Remarks. – Urnalana cristata, new species, shares with U.
granulimera Galil, 2005a, a granulate dorsal surface ofcheliped merus, a medially carinate carapace withtumescences on hepatic and branchial regions, and carinate
palms. However, it differs from the latter in having
prominently carinate pereiopods and in the curved andsigmoid form of the apical process of the first male pleopod(cf. Galil, 2005a, for U. granulimera).

Distribution. – Australia, Japan, China, Taiwan, Vietnam,
Indonesia, the Philippines, Andaman Sea, Laccadive Sea(Galil, 2003a).

Etymology. – crista Latin, “crest, carina”, for the boldly
carinate palms and pereiopods.

Urnalana cristata, new species(Figs. 3D, 5J–L)

Distribution. – Known only from Panglao Island.
Material examined. – Holotype, male (CL 11.1 mm) (NMCR),
station T29, Biking, Panglao, 09°34.5'N 123°50.6'E, 77–84 m,
sponges, 1 Jul.2004. Paratype: 1 male, parasitized (CL 11.3 mm)(ZRC 2007.0623), same data as holotype.

Urnalana cumingii (Bell, 1855)(Fig. 3C)

Description. – Carapace subpentagonal, globose; regions ofcarapace indistinct. Dorsal surface of carapace minutely
punctuate, obsoletely carinate medially, mostly glabrous,
short setae near posterolateral margin. Front produced,
prominent, frontal margin unidentate, deflexed, postfrontal
region laterally concave. Antennular fossa sealed by basal
antennular segment. Outer orbital margin unisutured, anterior
margin of efferent branchial channel forms part of lower
orbital margin. External maxillipeds lacking setose fringe

Leucosia cumingii White, 1847: 48 (nomen nudum); Bell, 1855b:
290, Pl. 31 Fig. 3.
Leucosia cumingi, Estampador, 1937: 511.
Leucosia galantua, Tan, 1996: 1035, Figs. 4d, 5a.
Urnalana cumingii, Galil, 2005a: 15, Figs. 1c, 4c.
Material examined. – 1 male (CL 9.4 mm) (ZRC 2007.0624), station
S38, channel between Tagbilaran town (Bohol) and Panglao,
09°38.1'N 123°51.4'E, 3–3.5 m, sandy bottom with little mud, 30
May 2004.

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Colour. – Young specimen. Carapace whitish marbled withochre, a pair of rust-coloured spots on posterolateral margin
above first pereiopods. Palm with small rust coloured spot
medially on upper margin, fingers with rust stripe medially.
Pereiopods white with rust coloured spots, dactyls purple (Fig.
3C).

Urnalana elata (A. Milne-Edwards, 1874)
Leucosia elata A. Milne-Edwards, 1874: 41, Pl. 2 Fig. 2.
Urnalana elata, Galil, 2005a: 16, Figs. 1D, 5A.
Material examined. – 1 ovigerous female (CL 9.7 mm) (ZRC
2007.0625), station M7, Momo Beach, Panglao, 09°36.1'N
123°45.2'E, 0–3 m, coral reef and sea-grass, 1 Jun.2004; 1 female(CL 6.3 mm) (MNHN), station S42, Pamilacan Island, 09°30.1'N
123°55.5'E, 15–20 m, sand with this hard layer, 1 Jul.2004.

Distribution. – Australia, Papua New Guinea, Indonesia, thePhilippines (Galil, 2005a).

Fig. 5. Gonopods of: A, B, Alox bothros, new species, holotype male (CL 9.0 mm) (NMCR); C–G, Alox somphos, paratype male (CL 8.7mm) (ZMA 100.605); H, I, Alox chaunos, new species, holotype male (CL 6.8 mm) (NMCR); J–L, Urnalana cristata, new species, holotype
male (CL 11.1 mm) (NMCR). A, B, H–L, G1s; C, D, G1 with distal part broken off; E, distal part of G1 showing showing breakage point
and remaining part of structure; F, G2; G, male abdomen.

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Galil & Ng: Some Leucosiidae from the Philippines
Distribution. – Fiji, Australia, Papua New Guinea, Indonesia,
Tonkin Bay, South China Sea, India, Andamans, South
Africa, Madagascar, Comoro Islands, Gulf of Aden, Red Sea(Galil, 2005a), the Philippines (new record).

Distribution. – Marshall Islands, Samoa, New Caledonia,
Australia, Papua New Guinea, Japan, Indonesia, Comoro
Islands (Galil, 2005a), the Philippines (new record).

Urnalana foresti (Chen, 1989)
Urnalana whitei (Bell, 1855)(Fig. 3F)

Leucosia foresti Chen, 1989: 240, Fig. 28, pl. 1(10).
Material examined. – 1 males (CL 9.3 mm) (ZRC 2007.0626),
station T11, Maribohoc Bay, Bohol, 09°40.9'N 123°50.0'E, 78–95
m, sponges and muddy sand, 16 Jun.2004.

Leucosia whitei Bell, 1855a: 362.
Urnalana whitei, Galil, 2005a: 32, Figs. 3d, 9b. (see Galil, 2005a:
32 for synonymy)

Remarks. – This species was not covered in Galil’s (2005a)
revision of the genus but it clearly belongs there and is referredhere for the first time. Looking at the descriptions of Leucosia
foresti Chen, 1989, and L. minuta Chen & Xu, 1991, we
believe the two species are very close, although Chen & Xu
(1991: 61, fig. 10) and Chen & Sun (2002 : 450, Fig. 204)
argue that there are differences in the form of the carapace
and thoracic sinus structures. As both were described by thelate H.-L. Chen within two years of each other, are fromdifferent localities (L. minuta: Nansha Islands = Spratlys, L.
foresti: the Philippines), and we have not examined the types,
it seems best to keep them separate for the time being.

Material examined. – 1 male (CL 8.0 mm) (ZRC 2007.0629), station
T6, west of Baclayon, Bohol, 09°35.1'N 123°51.2'E, 34–82 m,sandy-mud, large sponges, 2 Jun.2004.

Colour. – Carapace brownish, paler posteriorly, dull reddish
triangular patch on cardiac region; cheliped merus duncoloured, anteromedian granules bright orange; carpus,
propodus with red patch; fingers bright red proximally (Fig.
3F).
Distribution. – Australia, Indonesia (Galil, 2005a), thePhilippines (new record).

Distribution. – The Philippines (Chen, 1989).
Urashima pustuloides (Sakai, 1961)
Randallia pustuloides, Sakai, 1961: 135, Pl. 3 Fig. 4; Chen, 1989:
219, Fig. 16; Tan, 1996: 1023, 1054.
Urashima pustuloides, Galil, 2003a: 417, Figs. 2f, 5g, h; Komatsu
et al., 2005: 106. (see Galil, 2003a : 417 for synonymy)

Urnalana margaritata (A. Milne-Edwards, 1874)
Leucosia margaritata A. Milne-Edwards, 1874: 42, Pl. 2 Fig. 3.
Urnalana margaritata, Galil, 2005a: 25, Figs. 2e, 7b.

Material examined. – Balicasag Island, tangle nets: 1 male (CL36.5 mm) (ZRC 2001.0372), 200–300 m, Dec.2000; 1 female (CL36.3 mm) (ZRC 2007.0630), 200–300 m, Jun.2002; 2 males (CL35.5, 37.0 mm) (ZRC 2001.0568), 50–500 m, 28 Nov.2002; 1
juvenile male (CL 25.8 mm) (ZRC 2007.0631), Dec.2003; 1 male
(CL 36.8 mm), 2 females (CL 37.9, 26.9 mm) (NMCR), 28
May.2004; 2 males (CL 35.1, 34.3 mm), 1 female (CL 26.8 mm)(ZRC 2007.0632), Maribohoc Bay, Bohol, 100–300 m, tangle nets,
Nov.2003–Apr.2004.

Material examined. – 2 ovigerous females (CL 6.5, 7.0 mm) (ZRC
2007.0627), station T32, Baclayon, Bohol, 09°36.4'N 123°53.8'E,
60–62 m, muddy-sand, 3 Jul.2004; 1 female (CL 6.0 mm) (NMCR),
station S25, Ubajan, Bohol, 09°41.5'N 123°51.0'E, 21 m, 23
Jul.2004.

Distribution. – New Caledonia, Palau Islands, Australia,
Papua New Guinea, Indonesia, the Philippines (Galil, 2005a).

Distribution. – Australia, Japan, Taiwan, Indonesia, thePhilippines (Galil, 2003a).

Urnalana pulchella (Bell, 1855)(Fig. 3E)
Leucosia pulchella Bell, 1855a: 363.
Urnalana pulchella, Galil, 2005a: 29, Figs. 3A, 8A. (see Galil,
2005a: 29 for synonymy)

ACKNOWLEDGEMENTS
The study started as a collaborative study of the Philippine
crab fauna by the second author and Lawrence Liao
(University of San Carlos, Cebu City, Cebu, USC), and weare grateful for all the kind help he has rendered over theyears. Father Florante Camacho, President of the Holy Name
University in Bohol, was also instrumental in his support of
the work. The 2004 and 2005 expeditions were conducted asa joint exercise between the Musèum national d’Histoire
Naturelle (MNHN, Paris), University of San Carlos, Bureauof Fisheries and Research (BFAR, Manila), National Museum

Material examined. – 1 parasitized female (CL 8.1 mm) (MNHN),
station T5, west of Baclayon, Bohol, 09°35.3'N 123°52.2'E, 84–
86.9 m, sandy mud, 2 Jun.2004; 2 males (CL 8.4, 9.2 mm) (ZRC
2007.0628), station T28, Biking-Catamaran, Panglao, 09°35.0'N
123°51.4'E, 80 m, fine sand and mud, 1 Jul.2004; 1 female (CL 6.0mm) (NMCR), station T33, Baclayon, Bohol, 09°36.0'N 123°53.7'E,
67–74 m, sand, 3 Jul.2004.

Colour. – See Fig. 3E.

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THE RAFFLES BULLETIN OF ZOOLOGY 2007
Chen, H. & H. Sun, 2002. Arthropoda Crustacea. Brachyura. Marine
primitive crabs. Fauna Sinica. Invertebrata, 30, Science Press,
Beijing, 597 pp., 4 colour pls., 16 pls.

of the Philippines (NMP, Manila), and the National Universityof Singapore. We thank Philippe Bouchet (MNHN), Danilo
Largo (USC), Ludi Ludivena (BFAR) and Marivene Manuel(PNM) for all their kind help. Philippe Bouchet’s support
and persistence in realizing these very successful expeditions
has been vital in the project’s success. Bertrand Richer deForges (Institut de Recherche pour le Développement,
Noumea) was key in helping us obtain many of the deeper
water fauna, and sorting many of the small taxa in the field.
We also thank Tan Swee Hee, Lai Chiu Yun, Chan Tin Yam,
Lin Chia-Wei and Ristyo Rahayu for the many hours spentin the field sampling, sorting and photographing the thousandsof specimens during the 2004 and 2005 expeditions. We also
thank Hironori Komatsu and Tohru Naruse for carefully
reading the manuscript and suggesting many useful changes.
For the 2004 and 2005 expeditions, we acknowledge the kindfinancial assistance of the TOTAL Foundation, and the strong
support of Bernard Tramier. The study has been partially
supported by a research grant from the National Universityof Singapore. The first author thanks a Raffles Museum
research fellowship for funding her stay in Singapore, andthe entire laboratory personnel for making her stay in
Singapore memorable. Tohru Naruse, Hironori Komatsu andan anonymous reviewer kindly read and made many usefulcomments on the manuscript.

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