Ng et al.: Revision of Parilia
Taxonomy & Systematics
RAFFLES BULLETIN OF ZOOLOGY 66: 300–319

Date of publication: 4 May 2018
http://zoobank.org/urn:lsid:zoobank.org:pub:CBB28174-9B04-4A65-8C7C-5EB4B3608D98

The genus Parilia Wood-Mason, in Wood-Mason & Alcock, 1891, withdescription of a new species and establishment of a new genus for P.
tuberculata Sakai, 1961 (Crustacea, Brachyura, Leucosiidae)
Peter K. L. Ng1*, Suvarna Devi2 & Appukuttannair Biju Kumar2
Abstract. The taxonomy of the Indo-West Pacific leucosiid crabs of the genus Parilia Wood-Mason, in Wood-Mason& Alcock, 1891, is revised. The genus is redefined and four species are here recognised: P. alcocki Wood-Mason,in Wood-Mason & Alcock, 1891 (type species); P. major Sakai, 1961; P. ovata Chen, 1984; and P. pattersoni,
new species. Parilia pattersoni from the eastern Indian Ocean is morphologically similar to P. major but can easily
be distinguished by its more granulate carapace, chelipeds and ambulatory legs. Parilia tuberculata Sakai, 1961,
is referred to its own genus, Neparilia, new genus, as the structure of its carapace, third maxillipeds, chelipeds,
male pleon and male first gonopods are markedly different from Parilia s. str.
Key words. Crustacea, Leucosioidea, revision, systematics, new genus, new species, Indo-West Pacific

INTRODUCTION

genus. Specimens examined are deposited in the Department
of Aquatic Biology and Fisheries, University of Kerala(DABFUK), India; Centre of Advanced Study in Marine
Biology, Annamalai University (CASAU), Parangipettai,
Tamil Nadu, India; and the Zoological Reference Collection(ZRC) of the Lee Kong Chian Natural History Museum,
National University of Singapore. The morphological terms
used follow Ihle (1918) with changes as suggested by Ng
et al. (2008) and Davie et al. (2015). The measurements
provided (in millimetres) are of the maximum carapace
width and length, respectively.

The leucosiid genus Parilia Wood-Mason, in Wood-Mason& Alcock, 1891, was established with Parilia alcocki WoodMason, in Wood-Mason & Alcock, 1891, as the type species.
Three other species have since been added: P. major Sakai,
1961; P. tuberculata Sakai, 1961; and P. ovata Chen, 1984
(Ng et al., 2008). Parilia alcocki has a wide range in the
Indian Ocean (Prema et al., 2018), while P. major is widelydistributed in the western Pacific (Galil & Ng, 2015) with
two unconfirmed records from the eastern Indian Ocean (cf.
Sakai, 1961; Vidhya et al., 2017). Two other species havemore localised distributions: P. tuberculata is known only
from Japan (Sakai, 1976) and P. ovata has been reported anddescribed only from the South China Sea and Philippines(Tan, 1996).

SYSTEMATICSSuperfamily Leucosioidea Samouelle, 1819
Family Leucosiidae Samouelle, 1819

In this paper, we revise the taxonomy of Parilia. A new
species which superficially resembles P. major from theIndian Ocean is recognised and can be distinguished bypossessing a more pronounced granulation and spinulation
on its carapace and pereopods. Parilia tuberculata is shown
to be significantly different from other Parilia species in not
only possessing a strongly tuberculate carapace, but also in theform of its male thoracic sternum, male pleon, and male firstgonopod. It is here referred to its own genus, Neparilia, newParilia Wood-Mason, in Wood-Mason & Alcock, 1891
Type species. Parilia alcocki Wood-Mason, in Wood-Mason& Alcock, 1891, subsequent designation under Article 68.2.1[ICZN, 1999]; gender feminine).
Diagnosis. Carapace transversely subovate, ovate or
longitudinally ovate; dorsal surface covered with smallrounded granules or appearing smooth. Cardiac region withposteriorly directed spine; intestinal region with 2 spines.
Exopod of third maxilliped very broad, much wider thanischium and merus, with outer margin strongly convex;
coxa very wide, appearing almost semicircular, bracketingbase of exopod. Suborbital region prominently compressedby expanded buccal cavity. Adult male chelipeds elongate,
at least 3 times longer than carapace length; chela elongate,
fingers distinctly shorter than palm, in large male individuals,

Lee Kong Chian Natural History Museum, Faculty of Science, National University ofSingapore, 2 Conservatory Drive, Singapore 117377, Republic of Singapore; Email:peterng@nus.edu.sg (*corresponding author)
1

Department of Aquatic Biology & Fisheries, University of Kerala, Kariavattom,
Thiruvananthapuram – 695581, Kerala, India
2© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)

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fingers forming basal gape when closed. Male pleon withsomites 1 and 2 free, somites 3–5 completely fused, somite 6
and telson free; surface of somite 6 smooth; telson triangular,
subequal in length to somite 6. G1 elongate, slender, subdistal
part bifurcated, longer subdistal lateral projection directed
laterally inwards towards median of thoracic sternum;
projections flared, forming distinct serrulate or denticulate
flange. G2 distal segment as long as basal segment, withcurved flagelliform tip.

visible in dorsal view, frontal lobes distinctly triangular (Fig.
3A); exopod of third maxilliped in adult very broad, appears
foliaceous (Figs. 4B, 5A, C); surfaces of chelipeds andambulatory legs covered with small tubercles and granules(Figs. 1A, 2A, 6A); G1 very elongate, distal part prominently
flared, subdistal lateral projection dorso-ventrally flattened(Fig. 8A–C); female sternopleonal cavity with prominent
projection between vulvae (Fig. 9B). Indian Ocean.
Remarks. Prema et al. (2018) redescribed and figured thisspecies at length and there is no need to elaborate on its
taxonomy here. Parilia alcocki is distinct among congeners
in possessing a transversely ovate carapace which is very
swollen, and an exopod of the third maxilliped which is so
broad in adults that it appears foliaceous.

Remarks. Parilia Wood-Mason, in Wood-Mason & Alcock,
1891, is here restricted for three species, with P. tuberculata
Sakai, 1961, referred to its own genus (see remarks fornext genus).
Parilia alcocki and P. major are the two largest leucosioids
known in the world, with adults exceeding 70 mm incarapace width. Alcock (1899: 29) recorded males of P.
alcocki 56 mm in carapace width with a largest male in
the present study measuring 70.6 mm (CASAU). For P.
major, Chen & Sun (2002: 366) recorded a large male 60.1
mm in carapace width while in the present material, there
is a male measuring 70.0 mm (ZRC 2001.552). Myropsis
quinquespinosa Stimpson, 1871, from the western Atlantic
coast has been reported to grow to carapace widths of 65.0mm and 68.0 mm by Rathbun (1937: 166) and Williams(1984: 287), respectively.

Distribution. Known from the east coast of India and Bay
of Bengal (Wood-Mason & Alcock 1891; Wood-Mason,
1892; Alcock & Anderson, 1894; Alcock, 1896; Alcock,
1899; Huys et al., 2014; Krishnamoorthy, 2009; Lakshimi
Pillai et al., 2013; Prema et al., 2018), Arabian Sea (Lloyd,1907) including Pakistan (Tirmizi & Serène, 1971). The
species occurs at depths between 128–457 m (Alcock, 1896,
1899; Lloyd, 1907; Prema et al., 2018).
Parilia major Sakai, 1961
(Figs. 1B, 2B, 3B, 4C, D, 5D, E, 6B, 7C, D, 8E–I, 9C, D)
Parilia major Sakai, 1961: 137, pl. 3 fig. 5; Sakai, 1976: 105,
pl. 31 fig. 1 (Japan); Chen, 1989: 233, text fig. 24, pl. II, 1(Philippines); Chen, 1990: 180, pl. 3E; Takeda, 1993; Tan,
1996: 1046, fig. 7f–k; Ikeda, 1998: 86, 87 (Japan); Jeng etal., 1998: 121; Tan et al., 2000: 143, fig. 1d; Ng et al., 2001:
9 (Taiwan); Chen & Sun, 2002: 365, text fig. 162, pl. III–3;
Ng et al., 2008: 92; Galil, 2007: 560; Galil & Ng, 2007: 87(Philippines); Galil, 2015: 4; Galil & Ng, 2015: 460, fig. 4B;
Prema et al., 2018: figs. 3, 4D, F, 5H–K.

Parilia alcocki Wood-Mason, in Wood-Mason &Alcock, 1891
(Figs. 1A, 2A, 3A, 4A, B, 5A–C, 6A, 7A, B, 8A–D, 9A, B)
Parilia alcocki Wood-Mason, in Wood-Mason & Alcock, 1891:
264; Wood-Mason, 1892: pl. 5 fig. 3, 3a; Alcock & Anderson,
1894: 177; Alcock, 1896: 198; Alcock, 1899: 28, pl. 4 fig. 1;
Tirmizi & Serène, 1971: 30, text fig. 4 fig. 1; Tirmizi & Serène,
1971: 30, text fig. 4; Ng et al., 2008: 92; Krishnamoorthy,
2009: 2; Huys et al., 2014: 16; Lakshimi Pillai et al., 2013:72, unnumbered colour figure; Prema et al., 2018: 112, figs.
1, 2, 4A–D, F, 5A–G.
Parilia (sic) alcocki – Lloyd, 1907: 3.

Material examined. 1 male, 2 females (ZRC 2001.50),
Su-Ao Port, Nangfangao, northeastern Taiwan, coll. P.K.L.
Ng, 5 November 2000; 1 male, 3 females (ZRC 2017.217),
station PN1, Balicasag, Panglao, Bohol, Philippines, coll.
P.K.L. Ng from fishermen, June 2004; 7 males (largest 70.0
× 69.0 mm), 4 females (largest 48.3 × 49.2 mm), 2 ovigerous
females (ZRC 2001.552), Panglao, Bohol, Philippines, coll.
fishermen, 28 November 2001; 10 males, 5 females (ZRC
2017.218), Maribojoc Bay, Panglao, Bohol, Philippines, coll.
J. Arbasto, November 2003 – April 2004. For additional
material from Philippines and Papua New Guinea, see Galil& Ng (2007, 2015).

Material examined. 14 males (largest 70.6 × 52.2 mm,
smallest 43.0 × 36.5 mm), 10 ovigerous females (largest
47.4 × 38.8 mm, smallest 41.2 × 34.4 mm) (CASAU), 1
male (66.1 × 57.4 mm), 2 ovigerous females (45.2 × 37.0mm, 41.5 × 35.6 mm) (ZRC 2017.215), Pazhayar fish
landing center, 11°21′30.28″N 79°49′39.83″E, Tamil Nadu,
southeast India, 549 m, coll. M. Prema, 14 March 2017; 1
young male (24.8 × 23.4 mm) (ZRC 2017.216), fishing port,
Tuticorin, Tamil Nadu, Southeast India, coll. A.B. Kumar
et al., March 2017.

Diagnosis. Carapace globose (Figs. 1B, 2B, 3B); dorsal
surface appears almost smooth or with very low granules(Fig. 3B); branchial regions not distinctly swollen laterally
and dorsally, lateral margin of carapace smooth (Fig.
3B); dome-shape in frontal view (Fig. 4C); frontal region
protruding anteriorly with buccal cavity and third maxillipeds
not visible in dorsal view, frontal lobes distinctly triangular(Fig. 3B); outer surface of third maxillipeds almost smoothor with low, flattened granules (Figs. 4C, D, 5D, E); exopodof third maxilliped in adult broad but not foliaceous (Fig.

Diagnosis. Carapace transversely subovate (Figs. 1A, 2A,3A); dorsal surface covered with numerous small granules(Fig. 3A); branchial regions prominently swollen laterally
and dorsally (Figs. 1A, 2A, 3A, 4A); surface of carapace,chelipeds and ambulatory legs smooth or covered withsmall tubercles and granules; adult cephalothorax box-likein frontal view (Figs. 1A, 2A, 3A, 4A, 6A); frontal region
set back such that part of buccal cavity and third maxillipeds301

Ng et al.: Revision of Parilia

Fig. 1. Overall view. A, Parilia alcocki Wood-Mason, in Wood-Mason & Alcock, 1891, male (66.1 × 57.4 mm) (ZRC 2017.218), India;
B, P. major Sakai, 1961, male (70.0 × 69.0 mm) (ZRC 2001.552), Philippines; C, P. ovata Chen, 1984, male (31.6 × 43.3 mm) (ZRC
2017.219), Philippines; D, P. pattersoni, new species, holotype male (53.0 × 52.6 mm) (DABFUK), India.

5D, E); surfaces of ambulatory legs and chelipeds completely
smooth (Figs. 1B, 2B, 6B); G1 very elongate, distal partslightly flared, subdistal lateral projection dorso-ventrally
flattened (Fig. 8E–H); female sternopleonal cavity without
projection between vulvae (Fig. 9D). Western Pacific.

Sun, 2002), Philippines (Chen, 1989; Tan, 1996; Galil &Ng, 2007) to Makassar (Indonesia) (Chen, 1990) Papua New
Guinea (Galil & Ng, 2015) and Solomon Islands (Galil,
2007). The species occurs at depths between 100–457 m(Alcock, 1899; Chen, 1989; Galil, 2015).

Remarks. Sakai (1961) described the species from threemale and two female specimens from Tosa Bay in Japan.
The depository for this material was not specified and it is
not known where they are now (see remarks for Neparilia
tuberculata below). The species, however, is very distinct andthere is now no doubt about its identity. Uncertainties about
the presence of P. major in the Indian Ocean by Sakai (1961)
and Tirmizi & Serène (1971) are clarified in this paper withthe discovery that this material actually belongs to a new
species, P. pattersoni (see remarks for this species below).

Parilia ovata Chen, 1984
(Figs. 1C, 3C, 4E, F, 5F, 6C, 7E, F, 8J–M)
Parilia ovata Chen, 1984: 482, figs. 1–6; Chen, 1989: 235, text
fig. 30d; Tan, 1996: 1048; Chen & Sun, 2002: 366, text fig.
163; Ng et al., 2008: 92.
Myra anomala Zarenkov, 1990: 62, fig. 5.

Distribution. Has been reported from Japan and the East
China Sea (Sakai, 1961, 1976; Takeda, 1993; Ikeda, 1998;
Marumura & Kosaka, 2003), mainland China (Chen & Sun,
2002), Taiwan (Tan et al., 2000; Ng et al., 2001; Chen &Diagnosis. Carapace longitudinally ovate (Figs. 1C, 3C);
dorsal surface smooth (Fig. 3C); branchial regions not
swollen laterally and dorsally, lateral margin of carapace

Material examined. 1 male (31.6 × 43.3 mm) (ZRC
2017.219), northwestern Panglao, Bohol, Philippines, coll.
J. Arbasto, 2006.

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Fig. 2. Overall view. A, Parilia alcocki Wood-Mason, in Wood-Mason & Alcock, 1891, female (45.2 × 37.0 mm) (DABFUK), India;
B, P. major Sakai, 1961, female (48.3 × 49.2 mm) (ZRC 2001.552), Philippines; C, P. pattersoni, new species, paratype female (58.2 ×
55.8 mm) (DABFUK), India.

Zarenkov (1990) described Myra anomala from one small
juvenile male 12.3 mm in carapace length from Tonkin Bay
in northern Vietnam at a depth of 180 m. Galil (2001: 414)
noted that this species “does not belong within Myra, as themale abdomen is depicted with an articulate sixth segment.”
All Myra species have male pleonal somite 6 completely
fused with somite 5 and their G1 structures are very different(Galil, 2001). Galil (2001), however, did not reassign Myraanomala to another genus. Ng et al. (2008) synonymised
Myra anomala Zarenkov, 1990, under Parilia ovata Chen,
1984, without comment. From Zarenkov’s (1990: 62, fig.
5-5–10) description and figures, Myra anomala is almost
certainly a junior synonym of P. ovata. The carapace ofM. anomala (cf. Zarenkov, 1990: fig. 5-5) resembles thatof P. ovata (Fig. 3C) except that the lateral tubercles are
relatively more prominent which is expected of a juvenile
specimen. The exopod of the third maxilliped of M. anomala(Zarenkov, 1990: fig. 5-9) is not as wide as that of theadult (Fig. 5F) but this has also been observed for juvenile
P. alcocki in which the structure is proportionately more
slender (see earlier). The male pleon of M. anomala hasentire, smooth (Fig. 3C); dome-shape in frontal view (Fig.
4E); frontal region protruding prominently anteriorly withbuccal cavity and third maxillipeds not visible in dorsal
view, frontal lobes rounded (Fig. 3C); cardiac spine very
long (Fig. 3C); outer surface of third maxillipeds smooth(Figs. 4E, F, 5F); exopod of third maxilliped in adultbroad but not foliaceous (Fig. 5F); surfaces of ambulatory
legs and chelipeds completely smooth (Figs. 1C, 6C); G1
proportionately shorter, distal part flared, subdistal lateralprojection large, very wide, fan-like, laterally flattened (Fig.
8J–L); female sternopleonal cavity condition not known.
South China Sea and Philippines.
Remarks. This is not a common species, with only a few
records. The original description was by Chen (1984) fromone male and one female specimens from 173 m in theSouth China Sea, with a subsequent record of a juvenile
from the Philippines by Tan (1996). We here confirm therecord from the Philippines with the discovery of an adultmale from Bohol.

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Fig. 3. Dorsal view of carapace. A, Parilia alcocki Wood-Mason, in Wood-Mason & Alcock, 1891, male (66.1 × 57.4 mm) (ZRC 2017.218),
India; B, P. major Sakai, 1961, male (70.0 × 69.0 mm) (ZRC 2001.552), Philippines; C, P. ovata Chen, 1984, male (31.6 × 43.3 mm)(ZRC 2017.219), Philippines; D, P. pattersoni, new species, holotype male (53.0 × 52.6 mm) (DABFUK), India.

Distribution. Parilia ovata was described from the South
China Sea (Chen, 1984; Chen & Sun, 2002) but has also
been reported from the Philippines (Tan, 1996) and northern
Vietnam (Zarenkov, 1990, as Myra anomala). The few
specimens of this species have been found at depths between
173–230 m (Chen, 1984; Zarenkov, 1990).

the shape of typical Parilia, with somite 6 and telson free
and the surfaces unarmed (Zarenkov, 1990: fig. 5-10); andthe fingers of the chela are shorter than the palm (Zarenkov,
1990: fig. 5-7). The G1 of M. anomala is that of a juvenile,
being not distinctly elongate, and the subdistal part appears
to be damaged, although it appears that there was a lateralprojection at that junction (Zarenkov, 1990: fig. 5-6).
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Fig. 4. Frontal view of cephalothorax and third maxillipeds. A, B, Parilia alcocki Wood-Mason, in Wood-Mason & Alcock, 1891, male
(66.1 × 57.4 mm) (ZRC 2017.218), India; C, D, P. major Sakai, 1961, male (70.0 × 69.0 mm) (ZRC 2001.552), Philippines; E, F, P.
ovata Chen, 1984, male (31.6 × 43.3 mm) (ZRC 2017.219), Philippines; G, H, P. pattersoni, new species, holotype male (53.0 × 52.6mm) (DABFUK), India.

Parilia pattersoni, new species(Figs. 1D, 2C, 3D, 4G, H, 5G–I, 6D, 7G, H, 8N–S, 9E, F)

Material examined. Holotype: male (53.0 × 52.6 mm)(DABFUK), Tuticorin fishing port, Tamil Nadu, southeastern
India, coll. A.B. Kumar et al., March 2017. Paratypes: 2
males (47.7 × 48.6 mm, 72.5 × 69.8 mm), same data as
holotype; 3 males (64.4 × 63.5 mm, 54.1 × 53.8 mm, 50.0
× 49.4 mm), 2 females (58.2 × 55.8 mm, 52.6 × 54.0 mm,
51.0× 51.7 mm, 46.0 × 45.7 mm) (DABFUK), same location
as holotype, coll. R. Ravinesh, February 2017.

Parilia alcocki – Doflein, 1904: 44, pl. 14 figs. 8, 9 (not WoodMason, in Wood-Mason & Alcock, 1891).
Parilia major – Vidhya et al., 2017: 2334 (not Sakai, 1961).

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Diagnosis. Carapace globose (Figs. 1D, 2C, 3D); dorsal
surface covered with numerous small granules (Fig. 3D);
branchial regions not distinctly swollen laterally and dorsally,
lateral margin of carapace covered with small granules (Fig.
3D); dome-shape in frontal view (Fig. 4G); frontal region
protruding anteriorly with buccal cavity and third maxillipeds
not visible in dorsal view, frontal lobes distinctly triangular(Fig. 3D); outer surface of third maxillipeds covered withnumerous small granules (Figs. 4G, H, 5G–I); exopod of
third maxilliped in adult broad but not foliaceous (Fig.
5G–I); surfaces of ambulatory legs and chelipeds covered
with numerous distinct small sharp tubercles and granules(Figs. 1D, 2C, 6D); G1 very elongate, distal part flared,
subdistal lateral projection dorso-ventrally flattened (Fig.
8N–R); female sternopleonal cavity without projection
between vulvae (Fig. 9F).

Buccal cavity wide, about half width of carapace; notprotruding anteriorly; third maxillipeds sloping in lateralview when closed (Fig. 4G, H). Third maxillipeds completely
covering buccal space when closed; surfaces with numerous
small but distinct granules; palp (carpus, propodus anddactylus) concealed behind merus when retracted, longer
than merus; merus shorter than ischium, with triangular distal
margin, lined with setae; ischium with very shallow submedianoblique sulcus; exopod very broad but not foliaceous, wider
than ischium with outer margin strongly convex; main partof coxa deeply inserted into cephalothorax, external surface
with wide, almost semicircular plate extending from base,bracketing base of wide exopod (Figs. 4H, 5G–I). Surfacesof sub-orbital, pterygostomial, sub-hepatic and sub-branchialregions covered with distinct small granules; anterior
margin of pterygostomial region subcristate; sub-branchial
and ptergygostomial regions separated by distinct obliquegroove (Fig. 4G, H).

Description of adult male. Carapace almost globose,slightly longer than broad; surface covered with numerous
small but clearly visible granules (more prominent inlarge specimens); hepatic region adjacent to anterolateral
margin not prominently swollen, just visible in dorsal view;
gastric and branchial regions convex but not prominently
swollen; dorsal surface evenly convex; oblique cervicalgroove very shallow, not clearly visible, with a shallow
pit marking junction with gastric region; cardiac region
swollen, with posteriorly directed tooth, separated fromgastric and branchial regions by distinct sub-longitudinalgrooves; gastro-cardiac groove distinct, separating cardiac
and intestinal regions from swollen branchial regions, small
rugose area present at junction of groove between anterior
part of cardiac region and branchial region; low but visible
median longitudinal ridge present from gastric to cardiac
region; intestinal region low, pushed behind by swollen
cardiac region, not well defined; carapace appears dome-likefrom frontal view (Figs. 1D, 3D, 4G). Antennae with shortflagellum, not extending beyond orbit (Fig. 4G). Antennules
folding obliquely (Fig. 4G). Orbit small, rounded; eye withshort peduncle; supraorbital margin with 2 prominent fissures,
inner one longer; suborbital margin with distinct low toothon inner margin (Fig. 4G, H). Epistome wide, median partnot projecting anteriorly, with deep median fissure; lateralpart of epistome not visible from dorsal view; endostomial
ridge strong, longitudinal (Fig. 4G, H).

Chelipeds very long, slender, surfaces of merus, carpus andpalm covered with numerous small distinct sharp granules;
merus long, cylindrical; carpus short; palm subcylindrical,
with distal part proportionately broader, elongated, at least
twice length of carapace in largest specimens; palm more than
2.5 times length of slender fingers; dactylus gently curved;
basal part of propodal finger slightly arched, forming justvisible gape with dactylus when closed; cutting edges offingers (occluding surfaces) with small teeth (Figs. 1D, 6D).
Ambulatory legs long, slender; first leg longest, last leg
shortest; surfaces of merus, carpus and propodus covered withsmall sharp granules; merus and propodus subcylindrical, notarmed with spines or setae; dactylus gently curved, dorsoventrally flattened, each dorsal margin with dense row ofsetae, tip rounded (Fig. 1D).
Thoracic sternum with surfaces almost smooth; sternites
1–4 completely fused, forming one plate without trace of
sutures (Fig. 7G, H); sutures between sternites 4–8 mediallyinterrupted; pleonal locking mechanism consisting of
projecting edges of sternites 5–7, no trace of peg or tubercle
on sternite 5. Sternopleonal cavity deep, reaching to justbefore groove separating thoracic sternites 2 and 3 (Fig.
7G). Small part of thoracic 8 exposed when pleon closed,
visible as small plate at anterolateral edge of somite 3 (Fig.
7H). Penis tubular, exiting on condyle of coxa of fourth
ambulatory leg.

Frontal margin distinct, produced anteriorly, bilobed, withbroadly triangular concavity separating broadly triangularlobes; most of ptergygostomial regions and epistome notvisible in dorsal view (Figs. 1D, 3D). Anterior half ofanterolateral margin relatively low, starting as low ridgeof granules from supraorbital margin, becoming more
prominently cristate and granular towards posterior half whichhas 2 or 3 low teeth, the last one most prominent (Figs. 1D,
3D). Pterygostomial region wide, not visible in dorsal view;
anterior tooth adjacent to buccal cavity bilobed, separated
from margin by cleft; posterior tooth before anterolateral
margin low but distinct (Fig. 4G). Posterolateral margin
strongly convex, entire except for tooth just before posterior
carapace margin (Figs. 1D, 3D). Posterior carapace margin
with 2 prominent lateral sharp teeth (Figs. 1D, 3D).

Pleon elongate, triangular, surface smooth; somites 1 and 2
free; somites 3–5 completely fused, with only short transverse
median groove visible between somites 3 and 4, lateral
margin sinuous; somite 3 widest, lateral margins reachingcoxae of fourth ambulatory legs; somite 6 long, trapezoidal;
telson acutely triangular (Fig. 7G, H). G1 elongate, basal
part broadest, median and distal parts slender, tubular, gently
curved from ventral view; distal part bifurcated, longer
projection directed inwards towards median of thoracic
sternum, distal part flared to form prominent dorso-ventrally
flattened flange, margin of flange serrulate, shorter projection
in line with rest of G1, distal part flared with denticulate306

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Fig. 5. Right third maxilliped. A, Parilia alcocki Wood-Mason, in Wood-Mason & Alcock, 1891, male (66.1 × 57.4 mm) (ZRC 2017.218),
India; B, P. alcocki Wood-Mason, in Wood-Mason & Alcock, 1891, young male (24.8 × 23.4 mm) (ZRC 2017.216), India; C, P. alcocki
Wood-Mason, in Wood-Mason & Alcock, 1891, female (45.2 × 37.0 mm) (DABFUK), India; D, P. major Sakai, 1961, male (70.0 × 69.0mm) (ZRC 2001.552), Philippines; E, P. major Sakai, 1961, female (48.3 × 49.2 mm) (ZRC 2001.552), Philippines; F, P. ovata Chen,
1984, male (31.6 × 43.3 mm) (ZRC 2017.219), Philippines; G, P. pattersoni, new species, holotype male (53.0 × 52.6 mm) (DABFUK),
India; H, P. pattersoni, new species, paratype male (72.5 × 69.8 mm) (DABFUK), India; I, P. pattersoni, new species, paratype female(58.2 × 55.8 mm) (DABFUK), India.

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Fig. 6. Outer view of right chela. A, Parilia alcocki Wood-Mason, in Wood-Mason & Alcock, 1891, male (66.1 × 57.4 mm) (ZRC
2017.218), India; B, P. major Sakai, 1961, male (70.0 × 69.0 mm) (ZRC 2001.552), Philippines; C, P. ovata Chen, 1984, male (31.6 ×
43.3 mm) (ZRC 2017.219), Philippines; D, P. pattersoni, new species, holotype male (53.0 × 52.6 mm) (DABFUK), India.

Etymology. The species is named after Edward J. K.
Patterson, noted marine biologist and director of the Suganthi
Devadason Marine Research Institute in Tuticorin, Tamil
Nadu, India, who kindly hosted the authors during theirwork there.

margin on flange (Fig. 8N–R). G2 short, distal segment aslong as basal segment, with curved flagelliform tip (Fig. 8S).
Female. Chelipeds relatively shorter (Fig. 2C). Pleon strongly
convex, dome-like; somite 1 free, somites 2–6 functionally
fused with sutures still visible, telson mobile; telson
triangular with distinctly concave lateral margins; surface ofsomites smooth (Fig. 9E). Sternopleonal cavity large, deep,
with prominent rim, completely covered by pleon; sutures
separating thoracic sternites 4–8 all medially interrupted,
median part of cavity smooth; vulvae on anterior edge ofsternite 6, pushing into sternite 5, opening crescent-shaped,with narrow operculum, inner margin slightly raised (Fig.
9F). Eggs small, numerous, brown when fresh.

Remarks. Parilia pattersoni, new species, is morphologicallyclosest to P. major but can easily be distinguished by having
the surfaces of the ambulatory meri and chelipeds covered
with distinct small granules and sharp tubercles (Figs. 1D,
2C, 6D) (vs. completely smooth in P. major; Figs. 1B, 2B,
6B); the dorsal carapace surface and margins are covered withsmall sharp granules (Figs. 3D, 4G) (vs. smooth to almost
smooth with very small flattened granules in P. major; Figs.
3B, 4C); the surface of the pterygostomial and sub-branchialregions as well as the outer surface of third maxillipeds arecovered with distinct small granules (Figs. 4G, H, 5G–I) (vs.

Colour. Freshly caught, the adult carapace and chelipeds
are pale whitish-orange with the ambulatory legs white.
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Fig. 7. Male thoracic sternum and pleon. A, B, Parilia alcocki Wood-Mason, in Wood-Mason & Alcock, 1891, male (66.1 × 57.4 mm)(ZRC 2017.218), India; C, D, P. major Sakai, 1961, male (70.0 × 69.0 mm) (ZRC 2001.552), Philippines; E, F, P. ovata Chen, 1984,
male (31.6 × 43.3 mm) (ZRC 2017.219), Philippines; G, H, P. pattersoni, new species, holotype male (53.0 × 52.6 mm) (DABFUK), India.

Distribution. Known thus far only from eastern India (present
study) and western Sumatra in Indonesia (Doflein, 1904, as“P. alcocki”). Doflein’s (1904) specimen was from a depthof 141 m, and the present material was probably obtained
from depths between 100–300 m.

smooth or with very small, almost flattened granules in P.
major; Figs. 4C, D, 5D, E); the outer surface of the chela inboth sexes are distinctly covered with sharp granules (Fig. 6D)(vs. outer surface of chela completely smooth in P. major;
Fig. 6B); the cardiac spine is proportionately shorter evenin smaller specimens (Figs. 1D, 2C, 3D) (vs. spine always
proportionately longer in P. major; Figs. 1B, 2B, 3B); andthe lateral distal process of the G1 is relatively shorter (Fig.
8P, Q) (vs. relatively longer in P. major; Fig. 8F, G).

Key to Parilia species
1a. Carapace prominently elongated (Figs. 1C, 3C); cardiac spine
very long (Fig. 3C); lateral margin of carapace entire, smooth(Fig. 3C); frontal lobes rounded (Fig. 3C); lateral processes on
distal part of G1 very wide, fan-like (Fig. 8J–L) (South China
Sea, northern Vietnam and Philippines)....................................
......................................................... Parilia ovata Chen, 1984
1b. Carapace globose to subovate; cardiac spine long or short,never strongly elongated; lateral margin of carapace with 1–4
low tubercles, smooth or with small granules; frontal lobes
triangular; lateral processes on distal part of G1 flared but not
fan-like......................................................................................2

Doflein (1904: 44, pl. 14 figs. 8, 9) recorded and figured amale specimen (without chelipeds) of “Parilia alcocki” from
off the island of Nias in eastern Sumatra (incorrectly recorded
as a female) and there have been doubts about its precise
identity. Sakai (1961: 137) commented that the “P. alcocki
Doflein (nec Wood-Mason) may be a different species; thecarapace of Doflein’s species seems to be circular in outline
just as in the new species [P. major], but the merus andcarpus of the pereiopods are spinulated along the posteriorborder.” Tirmizi & Serène (1971: 31) agreed but deferredacting on it; neither did Prema et al. (2018). It is clear fromthe figure in Doflein (1904: pl. 14 figs. 8, 9) that the carapace
shape is quite different from that of P. alcocki s. str., being
globose rather than transversely subovate. Doflein’s (1904)
figures also show the carapace margins as lined with finegranules and the surfaces of the ambulatory meri covered
with spinules; characters associated with P. pattersoni, newspecies, as defined here. Vidhya et al. (2017) listed Parilia
major from the southeast coast of India but their record is
probably also P. pattersoni as well.

2a. Carapace transversely subovate, branchial regions prominently
swollen laterally and dorsally (Figs. 1A, 3A, 4A); box-like infrontal view (Fig. 4A); frontal region set back such that partof buccal cavity and third maxillipeds visible in dorsal view(Fig. 3A); exopod of third maxilliped in adult very broad,appears foliaceous (Figs. 4B, 5A, C); female sternopleonalcavity with prominent projection between vulvae (Fig. 9B)(Indian Ocean)................................................................ Parilia
alcocki Wood-Mason, in Wood-Mason & Alcock, 1891
2b. Carapace globose, branchial regions not distinctly swollen
laterally and dorsally (Fig. 3B, D); dome-shape in frontal
view (Fig. 4C, E); frontal region protruding anteriorly withbuccal cavity and third maxillipeds not visible in dorsal view309

Ng et al.: Revision of Parilia

Fig. 8. Gonopods. A–D, Parilia alcocki Wood-Mason, in Wood-Mason & Alcock, 1891, male (66.1 × 57.4 mm) (ZRC 2017.218), India;
E–I, P. major Sakai, 1961, male (70.0 × 69.0 mm) (ZRC 2001.552), Philippines; J–M, P. ovata Chen, 1984, male (31.6 × 43.3 mm)(ZRC 2017.219), Philippines; N–S, P. pattersoni, new species, holotype male (53.0 × 52.6 mm) (DABFUK), India. A, E, J, N, left G1(ventral view); O, left G1 (dorsal view); B, F, K, P, distal part of left G1 (ventral view); C, G, H, L, Q, R, distal part of left G1 (dorsalview); D, I, M, S, left G2. Scales: A, D, E, I, J, M, N, O, S = 2.0 mm; B, C, F–H, K, L, P–R = 0.5 mm.

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Fig. 9. Female pleon and sternopleonal cavity. A, B, Parilia alcocki Wood-Mason, in Wood-Mason & Alcock, 1891, female (45.2 × 37.0mm) (DABFUK), India; C, D, P. major Sakai, 1961, female (48.3 × 49.2 mm) (ZRC 2001.552), Philippines; E, F, P. pattersoni, newspecies, paratype female (58.2 × 55.8 mm) (DABFUK), India(Fig. 3B, D); exopod of third maxilliped in adult broad but
not foliaceous (Fig. 5D, E, G–I); female sternopleonal cavity
without projection between vulvae (Fig. 9D, F).....................3

4H, 5G–I); surface of ambulatory legs, chelipeds and chela
with numerous distinct small sharp tubercles and granules(Figs. 1D, 2C, 6D) (eastern Indian Ocean)...............................
................................................. Parilia pattersoni, new species

3a. Lateral margin of carapace smooth (Fig. 3B); outer surface ofthird maxillipeds smooth or with low, flattened granules (Figs.
4D, 5D, E); surface of ambulatory legs, chelipeds and chela
completely smooth (Figs. 1B, 2B, 6B) (western Pacific) ........
......................................................... Parilia major Sakai, 1961
3b. Lateral margin of carapace lined with small granules (Fig. 3D);
outer surface of third maxillipeds distinctly granulated (Figs.

Neparilia, new genus
Diagnosis. Carapace ovate; dorsal surface covered large
rounded tubercles and granules. Cardiac region with low
rounded tubercle; intestinal region with 2 short spines.
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Fig. 10. Neparilia tuberculata (Sakai, 1961), male (37.0 × 34.4 mm) (ZRC 2017.1052), Philippines. A, overall view; B, dorsal view ofcarapace; C, right third maxilliped.

Exopod of third maxilliped broad, distinctly wider thanischium and merus, with outer margin strongly convex;
coxa wide, not semicircular in shape, not bracketing baseof exopod. Suborbital region prominently compressedby expanded buccal cavity. Adult male chelipeds about
2 times longer than carapace length; chela not elongate,
fingers longer than palm, in large male individuals, closingtightly. Male pleon with somites 1 and 2 free, somites 3–5completely fused, somite 6 separated from somite 5 by suture
but immobile, telson free; distal and proximal surfaces ofsomite 6 each with a distinct cup-shaped tubercle, that on
distal part larger; telson elongate, linguiform, longer thansomite 6. G1 slender, distal part tapering without lateralprojection or flanges. G2 distal segment as long as basal
segment, with curved flagelliform tip.

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Fig. 11. Neparilia tuberculata (Sakai, 1961), male (37.0 × 34.4 mm) (ZRC 2017.1052), Philippines. A, B, frontal view of cephalothorax
and third maxillipeds; C, D, thoracic sternum and pleon; E, pleon; F, pleonal somite 6 showing cup-like tubercles on distal and proximal
surfaces; G, outer view of right chela.

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Fig. 12. Neparilia tuberculata (Sakai, 1961), male (37.0 × 34.4 mm) (ZRC 2017.1052), Philippines. A, left G1 (ventral view); B, distal
part of left G1 (ventral view); C, distal part of left G1 (dorsal view); D, left G2. Scales: A, D = 1.0 mm; B, C = 0.5 mm.

Type species. Parilia tuberculata Sakai, 1961, by monotypy
and present designation.
Etymology. From the Latin “ne-” for not, alluding to thepresent realisation that the type species is not a member ofParilia. Gender feminine.

6 in Parilia; Fig. 7A, C, E, G); and the G1 is straight,
without a subdistal lateral projection, not flared and lacks
serrulate flanges distally (Fig. 12A–C) (vs. distal part of G1
bifurcated, with prominent lateral projection, each projection
fared with serrulate or denticulate flanges in Parilia; Fig.
8A–C, E–H, J–L, N–R).

Remarks. Sakai (1961) placed this species in Parilia because
it possessed a very broad exopod of the third maxilliped. Itis, however, not possible to retain P. tuberculata in Parilia
because the coxa of the third maxilliped is not substantiallylaterally expanded or semicircular (Figs. 10C, 11B) (vs. coxa
prominently expanded laterally, appearing semicircular in
Parilia; Figs. 4, 5); the adult male chelipeds are relatively
short (about 2 times carapace length), the chela are notelongate with the fingers longer than the palm (Figs. 10A,
11G) (vs. adult male chelipeds more than 3 times longer
than carapace length, chela prominently elongate with fingers
much shorter than palm in Parilia; Figs. 1, 6); the surfacesof the ambulatory meri are covered with short, sharp spines(Figs. 10A, 11C, D) (vs. surface of ambulatory meri smoothor covered with granules, never spiniform in Parilia; Figs.
1, 2); the male pleonal somites 5 and 6 are functionally
fused even though there is a distinct suture separating themand somite 6 has a cup-like tubercle on both the distal andproximal surfaces, with the distal one more prominent (Fig.
11C, E, F) (vs. male pleonal somites 5 and 6 mobile, surface
of somite 6 smooth in Parilia; Fig. 7A, C, E, G); the maletelson is linguiform and much longer than somite 6 (Fig.
11C, E) (vs. telson triangular, subequal in length to somite

The carapace and general external features of Neparilia
resemble Galilia Ng & Richer de Forges, 2007 (type species
Galilia narusei Ng & Richer de Forges, 2007), but several
features easily distinguish them. In Galilia, the exopod of thethird maxilliped is transversely narrower than the endopod(Ng & Richer de Forges, 2007: fig. 2E; Komai & Tsuchida,
2014: figs. 19B, 21A) (vs. exopod is distinctly broader thanthe endopod in Neparilia; Figs. 10C, 11B); the intestinal
region has two large rounded tubercles (Ng & Richer deForges, 2007: fig. 2F; Komai & Tsuchida, 2014: fig. 19A,C) (vs. intestinal region with two short spines in Neparilia;
Figs. 10B, 11D); the fingers of the chela are shorter thanthe palm (Ng & Richer de Forges, 2007: fig. 3D–F; Komai& Tsuchida, 2014: fig. 20A) (vs. fingers distinctly longer
than the palm in Neparilia; Fig. 11G); the male telson isprominently elongate with the male sternopleonal cavity
reaching to the suture between thoracic sternites 2 and 3 (Ng& Richer de Forges, 2007: fig. 3A, B; Komai & Tsuchida,
2014: figs. 19F, 21B) (vs. male telson proportionately
shorter and the male sternopleonal cavity reaches only tothe midpoint of sternite 3 in Neparilia; Fig. 11C, E); thesurface of male pleonal somite 6 has a truncate tubercle on
the distal margin (Ng & Richer de Forges, 2007: fig. 3A,
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Fig. 13. Freshly collected male (not available for examination) from southern Philippines by LUMIWAN 2008 cruise. A, overall view;
B, C, frontal view of cephalothorax D, ventral view showing third maxillipeds and anterior thoracic sternum. Photographs by T.-Y. Chan.

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Fig. 14. Urashima pustuloides (Sakai, 1961), male (44.5 × 37.6 mm) (ZRC 2017.1069), Philippines. A, overall view; B, C, frontal viewof cephalothorax and third maxillipeds; D, right third maxilliped; E, anterior thoracic sternum, telson and pleonal somites 5 and 6 showing
spine on distal margin of somite 6; F, thoracic sternum and pleon; G, outer view of right chela.

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Fig. 15. Urashima pustuloides (Sakai, 1961), male (44.5 × 37.6 mm) (ZRC 2017.1069), Philippines. A, left G1 (ventral view); B, distal
chitinous part of left G1 (ventral view); C, distal chitinous part of left G1 (subventral view); D, distal chitinous part of left G1 (dorsalview); E, left G2. Scales: A, E = 2.0 mm; B–D = 0.5 mm.

Neparilia tuberculata (Sakai, 1961)(Figs. 10–13)

B; Komai & Tsuchida, 2014: fig. 19F) (vs. the surface has
a cup-like tubercle on both the proximal and distal surfaces
in Neparilia; Fig. 11C, E); male pleonal somites 5 and 6
are completely fused, separated at most by a shallow suture(Ng & Richer de Forges, 2007: fig. 3A; Komai & Tsuchida,
2014: fig. 19F) (vs. somites 5 and 6 are free and separated
by a deep suture in Neparilia; Fig. 11C, E); and the distal
third of the G1 is prominently bent (Ng & Richer de Forges,
2007: fig. 4G, H; Komai & Tsuchida, 2014: fig. 21C, D;
Shih et al., 2013: fig. 3A, B) (vs. G1 straight in Neparilia;
Fig. 12A–C).

Parilia tuberculata Sakai, 1961: 137, pl. 3 fig. 6; Sakai,
1976: 106, text fig. 58, pl. 30 fig. 3; Ikeda, 1998: 85; Ng
et al., 2008: 92.
Material examined. 1 male (37.0 × 34.4 mm) (ZRC
2017.1052), Balut Island, Mindanao, 150 m, Philippines,
coll. tangle net, local fishermen, from dealers, 2017.
Comparative material. Urashima pustuloides (Sakai, 1961):
2 males (44.5 × 37.6 mm, 44.6 × 40.8 mm), 2 females (32.0
× 27.0 mm, 46.2 × 39.3 mm) (ZRC 2017.1069), Balicasag
Island, Panglao, Bohol, Philippines, from tangle nets, coll.
P.K.L. Ng, November 2004.

Neparilia also superficially resembles Urashima Galil, 2003
(type species Randallia pustuloides Sakai, 1961) (Fig. 14A),
but in Urashima, the third maxilliped has the exopod only
slightly broader than the exopod (Fig. 14C, D); the fingersof the chela are shorter than the palm (Fig. 14G; Sakai,
1961: pl. 3 fig. 4; Sakai, 1976: pl. 30 fig. 1; Galil, 2003:
fig. 2E, F; Tan et al., 2000: fig. 3b) (vs. fingers distinctly
longer than the palm in Neparilia; Fig. 11G); the surface ofmale pleonal somite 6 has a spine on the distal margin (Fig.
14E; Galil, 2003: 416); and male pleonal somites 5 and 6
are completely fused (Fig. 14F; Galil, 2003: 416). While theG1s of the two genera are superficially similar in generalshape, that of Urashima has the distal quarter distinctly
chitinised and spatuliform (Fig. 15A–D) (vs. the distal part
is not chitinised and conical in Neparilia; Fig. 12A–C).

Diagnosis. As for genus.
Remarks. Sakai (1961) described the species from two malesin Tosa Bay, Japan, but it is not known where the types are
now kept. They are not in the National Museum of Natureand Science, Tsukuba (formerly National Science Museum,
Tokyo). Ng et al. (2017) discussed the matter of Sakai’smaterial at length, with extant material in the Smithsonian
Institution, Naturmuseum Senckenberg, Kanagawa Museum
or lost.
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Ng et al.: Revision of Parilia
Fortunately, the male specimen on hand agrees in all aspects
with the description and figures in Sakai (1961, 1976) and
we have no doubt of its identity. During a research cruise(LUMIWAN 2008) to southwestern Philippines in 2008
jointly organised by the Bureau of Fisheries and Aquatic
Resources of the Philippines, Philippine National Museum,Musèum national d’Histoire Naturelle (Paris), Lee Kong
Chian Natural History Museum, and National Taiwan Ocean
University, a male specimen of Neparilia tuberculata was
obtained and photographed fresh (Fig. 13). It agrees with themale specimen on hand in all key aspects although its G1
was not photographed. Unfortunately, the specimens fromthis cruise are not available for study for the time being and
we are unable to add this to our material examined. The
colour of the fresh specimen, however, is striking, beinga bright orange on all the dorsal surfaces with the ventral
regions white. This agrees well with the Japanese female
figured in Ikeda (1998: 85).

Indian Marine Survey between 1890 and 1891: implications
for malacostracan nomenclature. Zootaxa, 3757(1): 1–78.
ICZN, International Commission on Zoological Nomenclature(1999) International Code of Zoological Nomenclature.
International Commission of Zoological Nomenclature.
Fourth Edition. Adopted by the XXI General Assembly of
the International Union of Biological Sciences. International
Trust for Zoological Nomenclature, in association with the
British Museum (Natural History), London, 338 pp.Ihle JEW(1918) Die Decapoda Brachyura der Siboga-Expedition. III.
Oxystomata: Calappidae, Leucosiidae, Raninidae. Siboga
Expeditie Monografie, 39b2: 159–322, figs. 78–148.
Ikeda H (1998) The Deep-Sea Crabs of Sagami Bay. Hayama
Shiosai Museum, Kanagawa, Japan, 180 pp.
Jeng M-S, Shao K -T, Tzeng C-S, Feng F-L & Wu S-H (1998)
Survey and monitoring of natural ecological resources in the
Northeast Coast National Scenic Area (IV). Northeast Coast
National Scenic Area Administration, Bureau of the Ministryof Transportation and Communications, Taipei, 182 pp. [In
Chinese]
Komai T & Tsuchida S (2014) Deep-sea decapod crustaceans(Caridea, Polychelida, Anomura and Brachyura) collectedfrom the Nikko Seamounts, Mariana Arc, using a remotely
operated vehicle “Hyper-Dolphin”. Zootaxa, 3764: 279–316.
Krishnamoorthy P (2009) Brachyuran crabs from the collectionsof Marine Biological Centre. Records of the Zoological Surveyof India, Occasional Paper, 304: 1–46.
Lakshimi Pillai S, Kizhakudanand JK & Thirumilu P (2013)
Potential ornamental marine brachyuran crabs available ofChennai. Fishing Chimes, 33(4): 70–73.
Lloyd RE (1907) Contributions to the fauna of the Arabian Sea,
with descriptions of new fishes and Crustacea. Records of the
Indian Museum, 1: 1–12.
Marumura M & Kosaka A (2003) Catalogue of the Brachyuran andAnomuran Crabs Donated by the Late Mr. Seiji Nagai to the
Wakayama Prefectural Museum of Natural History. Wakayama
Prefectural Museum of Natural History, Kainan, 73 pp.
Ng PKL, Guinot D & Davie PJF (2008) Systema Brachyurorum:
Part I. An annotated checklist of extant brachyuran crabs of
the world. Raffles Bulletin of Zoology, Supplement 17: 1–286.
Ng PKL, Komai T & Sato T (2017) On the trail of a Japanese “ghost
species” – the identity of Goniopugettia tanakae Sakai, 1986,
and the establishment of a new genus for Pugettia sagamiensis
Gordon, 1930 (Decapoda, Brachyura, Epialtidae). Crustacean
Research, 46: 133–152.
Ng PKL & Richer de Forges B (2007) A new genus and new
species of leucosiid crab from New Caledonia, with a noteon the validity of Tanaoa serenei (Richer de Forges, 1983)(Crustacea: Decapoda: Brachyura). Zootaxa, 1662: 15–24.
Ng PKL, Wang CH, Ho PH & Shih HT (2001) An annotatedchecklist of brachyuran crabs from Taiwan (Crustacea:
Decapoda). National Taiwan Museum Special Publication
Series, Number 11: 1–86, 8 colour pls.
Prema M, Ravichandran S & Ng PKL (2018) Redescription of
Parilia alcocki (Wood-Mason, in Wood-Mason & Alcock,1891) (Decapoda, Brachyura, Leucosiidae) from southeast
India. Zootaxa, 4378(1): 111–120.
Rathbun MJ (1937) The oxystomatous and allied crabs of America.
Bulletin of the U. S. National Museum, 166: i–vi, 1–278, pls.
1–86.
Sakai T (1961) New species of Japanese crabs from the collectionof His Majesty the Emperor of Japan. Crustaceana, 3(2):131–150, figs. 1–4, pls. 3, 4.
Sakai T (1976) Crabs of Japan and the Adjacent Seas. In three
volumes; English Text, pp. xxix+773 pp., Japanese text,
pp. 1–461, Plates volume, pp. 1–16, pls. 1–251. Kodansha

Distribution. Neparilia tuberculata is known only fromJapan (Sakai, 1961, 1976; Ikeda, 1998; Marumura & Kosaka,
2003) and the Philippines (present record).
ACKNOWLEDGEMENTS
The authors thank R. Ravinesh for his assistance during ourfield work. ABK thanks the University Grants Commission
CAS programme for research support. Thanks are due to J.C.
Mendoza for reminding the first author about the specimen
collected from the LUMIWAN 2008 cruise, and to Chan
Tin-Yam for the photographs. The first author is grateful to
Bella Galil, Hironori Komatsu and Tohru Naruse for theircomments on the genus and help with the present work.
LITERATURE CITED
PariliaInvestigator Galil BS (2001) A revision of Myra Leach,
1817 (Crustacea: Decapoda: Leucosioidea). Zoologische
Mededelingen, 75(24): 409–446, figs. 1–19.Galil BS (2003)
Four new genera of leucosiid crabs (Crustacea: Brachyura:
Leucosiidae) for three new species and nine species previously
in the genus Randallia Stimpson, 1857, with a redescription of
the type species, R. ornata (Randall, 1939). Proceedings of the
Biological Society of Washington, 116(2): 395–422.
Galil BS (2007) The deep-water Calappidae, Matutidae andLeucosiidae of the Solomon Islands, with a description of a
new species of Euclosia Galil, 2003 (Crustacea, Decapoda,
Brachyura). Zoosystema, 29(3): 555–563.
Galil BS & Ng PKL (2007) Leucosiid crabs from Panglao,
Philippines, with descriptions of three new species (Crustacea:
Decapoda: Brachyura). Raffles Bulletin of Zoology, Supplement
16: 79–94.
Galil BS (2015) On a collection of Leucosioidea (Crustacea,
Decapoda, Brachyura) from Papua New Guinea, with thedescription of a new species. European Journal of Taxonomy,
155: 1–10.
Galil BS & Ng PKL (2015) Leucosiid crabs from Papua New
Guinea, with descriptions of eight new species (Crustacea:
Decapoda: Brachyura). Zootaxa, 4027(4): 451–486.
Huys R, Low MEY, De Grave S, Ng PKL & Clark PF (2014) On
two reports associated with James Wood-Mason and Alfred
William Alcock published by the Indian Museum and the318

RAFFLES BULLETIN OF ZOOLOGY 2018
Ltd., Tokyo.Samouelle G (1819) The Entomologist’s UsefulCompendium, or an Introduction to the Knowledge of the British
Insects. London, Thomas Boys, Shih Y-J, Ho P-H, Chan T-Y& Naruse T (2013) New records of leucosiid crabs (Decapoda,
Brachyura) from Taiwan. Crustaceana, 86: 728–738.
Stimpson W (1871) Brachyura. In: Preliminary Report on the
Crustacea dredged in the Gulf Stream in the Straits of Florida,
by L. F. de Pourtales, Assist. U.S. Coast Survey. Bulletin of
the Museum of Comparative Zoology at Harvard College,2(2): 109–160.
Takeda M (1993) Catalogue of Crab Specimens (Crustacea:
Decapoda: Brachyura) (Part 1). National Science Museum,
Tokyo, 93 pp.
Tan CGS (1996) Leucosiidae of the Albatross expedition to thePhilippines, 1907–1910 (Crustacea: Brachyura: Decapoda).
Journal of Natural History, 30(7): 1021–1058, figs. 1–8.
Tan SH, Wu S-H & Huang J-F (2000) Seven new records ofmarine crabs from Taiwan (Crustacea: Decapoda: Brachyura:
Calappidae, Lencosiidae and Cancridae). In: Hwang J-S, Wang
C-H & Chan T-Y (eds.) Proceedings of the International
Symposium on Marine Biology in Taiwan - Crustacean andZooplankton Taxonomy, Ecology and Living Resources.
National Taiwan Museum Special Publication Series, No. 10:
141–147.

Tirmizi NM & Serène R (1971) The rediscovery of two speciesof crabs (Decapoda, Brachyura) with observations on threeother species from Pakistan. Crustaceana, 2: 21–32, pls. 1, 2.
Vidhya V, Jawahar P & Karuppasamy K (2017) Annotated checklist of the brachyuran crabs (Crustacea: Decapoda) from Gulf ofMannar region, south east coast of India. Journal of Entomology
and Zoology Studies, 5(6): 2331–2336.
Williams AB (1984) Shrimps, Lobsters, and Crabs of theAtlantic Coast of the Eastern United States, Maine to Florida.
Smithsonian Institution Press, 550 pp.
Wood-Mason J (1892) Illustrations of the Zoology of H.M. Indian
Marine Surveying Steamer Investigator, under the command of
Commander A. Carpenter, R.N., D.S.O., and of Commander
R.F. Hoskyn, R.N. Part 1. Crustaceans, Plates I–V. Publishedunder the Authority of the Director of the Royal Indian Marine.
Calcutta: Printed and sold by the Superintendent of Government
Printing. [see Clark & Crosnier, 1992, for dates of publication]
Wood-Mason J & Alcock A (1891) Natural History Notes from
H.M. Indian Marine Survey Steamer “Investigator”, Commander
R.F. Hoskyn, R.N., commanding. No. 21.On the Results of
the last Season’s Deepsea Dredging. Annals and Magazine
of Natural History, March 1891, (6)7(39): 258–272. [For an
elaboration on the authorship of the taxa described in this seriesof papers, see HUYS et al., 2014.]

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