RAFFLES BULLETIN OF ZOOLOGY 2017
Taxonomy & Systematics
RAFFLES BULLETIN OF ZOOLOGY 65: 243–268

Date of publication: 14 July 2017
http://zoobank.org/urn:lsid:zoobank.org:pub:CC246EF9-E704-4DDC-BD25-61B6102A382F

On a collection of Homolidae from the South China Sea, withdescriptions of two new species of Homologenus A. Milne-Edwards,
in Henderson, 1888, and the identities of Homologenus malayensis
Ihle, 1912, and Lamoha superciliosa (Wood-Mason, in Wood-Mason& Alcock, 1891)
Peter K. L. Ng1* and Bertrand Richer de Forges2
Abstract. Recent collections from southern Taiwan and the South China Sea obtained 10 species of homolid crabs,of which two species of Homologenus A. Milne-Edwards, in Henderson, 1888, are described as new. The taxonomyof the allied H. malayensis Ihle, 1912, from Indonesia and Papua New Guinea is also clarified. The identity of
Lamoha superciliosa (Wood-Mason, in Wood-Mason & Alcock, 1891) s. str. from the Indian Ocean is discussed
and specimens from East Asian seas that had been referred to this species are here shown to be conspecific with
L. longirostris (Chen, 1986) instead. The taxonomy of L. superciliosa and L. longirostris is treated.
Key words. Taxonomy, new species, South China Sea, deep sea, homolid crabs, Homologenus

INTRODUCTION

The taxonomy of these species is treated in this paper, with
the new species diagnosed and figured. The terminology
essentially follows that used by Guinot & Richer de Forges(1995), with some recent changes recommended by Davieet al. (2015). The authorship and year of publication for thevarious species described by Wood-Mason & Alcock follow
Huys et al. (2014). Measurements provided (in millimetres)
are of the maximum carapace length and width (including
spines), respectively. Specimens examined are deposited in
the Muséum national d’Histoire naturelle, Paris (MNHN);
Oxford University Museum of Natural History, United
Kingdom (OUMNH); The Naturalis Biodiversity Center(including the zoological collections of the Amsterdam
University), Leiden (NNM ZMA); U.S. National Museum of
Natural History, Smithsonian Institution, Washington D.C.
(USNM); Crustacean Collection of the National Taiwan
Ocean University, Keelung, Taiwan (NTOU); Instituteof Oceanology, Chinese Academy of Sciences, Qingdao(IOCAS); and Zoological Reference Collection of the Lee
Kong Chian Natural History Museum (formerly the Raffles
Museum of Biodiversity Research), National University ofSingapore (ZRC).

In late 2015, Chan Tin-Yam of the National Taiwan Ocean
University in Keelung, Taiwan, passed the authors aninteresting collection of homolid crabs he and his colleagues
obtained from several cruises in the South China Sea,
including around Tungsha [= Dongsha] Island. The material,
while not extensive, is nevertheless interesting as it contains
representatives of a number of rare species. Examination ofthis material led us to reappraise the taxonomy of Lamoha
superciliosa (Wood-Mason, in Wood-Mason & Alcock,1891) and L. longirostris (Chen, 1986), with material
previously assigned to the former species from Taiwan now
synonymised with L. longirostris. Lamoha superciliosa s. str.
is now restricted to the Indian Ocean. The collection of agood series of specimens of “Homologenus malayensis Ihle,
1912” from Taiwan and the South China Sea also prompted
us to re-examine the taxonomy of this and allied species.
Two new species of Homologenus are here recognised, H.
exilis n. sp. and H. brevipes n. sp. Homologenus malayensis
Ihle, 1912 s. str., is redescribed and figured from a goodseries of specimens from Papua New Guinea.

TAXONOMYFamily Homolidae De Haan, 1833

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

Homola Leach, 1816

c/o Muséum National d’Histoire Naturelle, Institut de Systématique, Évolution,
Biodiversité
2

Homola orientalis Henderson, 1888

ISYEB - UMR 7205 – CNRS, MNHN, UPMC, EPHE, Département Systématiqueet Evolution, 57 rue Cuvier, CP26, 75005 Paris, France; Email: b.richerdeforges@
gmail.com

Homola orientalis Henderson, 1888: 19, pl. 2, fig. 1, 1a (see Guinot& Richer de Forges, 1995: 331; Richer de Forges & Ng, 2007:
30; for complete synonymy)© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)243

Ng & Richer de Forges: Homolidae from the South China Sea
Material examined. 1 female (14.7 × 11.5 mm) (ZRC
2016.0553), station CP 4116, 20°02.32′N 114°10.22′E– 20°02.92′N 114°11.29′E, Taiwan, South China Sea,
262–298 m, trawl, coll. NANHAI 2014 Cruise, 11 January
2014. – 1 male (soft, 30.7 × 22.9 mm) (ZRC 2016.0554),
station CP 4159, northwest of Tungsha Island, 20°45.92′N
116°41.11′E – 20°47.62′N 116°42.34′E, Taiwan, South China
Sea, 190–221 m, dead coral substrate, coll. ZHONGSHA
2015 Cruise, 30 July 2015.
Remarks. The taxonomy of this wide-ranging Indo-West
Pacific taxon, which is likely to be a species-complex, hasbeen discussed at length by Guinot & Richer de Forges(1995) and Richer de Forges & Ng (2007).
Lamoha Ng, 1998
Hypsophrys Wood-Mason, in Wood-Mason & Alcock, 1891
Lamoha Ng, 1998: 121 (replacement name for Hypsophrys WoodMason, in Wood-Mason & Alcock, 1891, preoccupied by
Hypsophrys Agassiz, 1859 [freshwater fish])

Lamoha superciliosa(Wood-Mason, in Wood-Mason & Alcock, 1891)(Figs. 1, 2, 5E, F, 6H, 7G, H)
Hypsophrys superciliosa Wood-Mason, in Wood-Mason &Alcock, 1891: 269. – Alcock & Anderson, 1895: pl. 14 fig.
4, 4a. – Alcock, 1900: 14; 1901: 67, pl. 6 fig. 24. – Bouvier,
1896: 32. – Van Straelen: 1928: 19. – Gordon, 1950: 220,
224. – Williams, 1974: 485. – Williams, 1976: 889. – Guinot& Richer de Forges, 1981: 543. – Guinot & Richer de Forges,
1995: 445, figs. 56c–e, 61i. – Ng, 1998: 121. – Ng et al., 2008:
40. – Huys et al., 2014: 27.

Fig. 1. Lamoha superciliosa (Wood-Mason, in Wood-Mason &Alcock, 1891). A, B, female (after Alcock, 1901: pl. 6 fig. 24);
C, D, male (after Alcock & Anderson, 1895: pl. 14 fig. 4, 4a).

The OUMNH specimen agrees well with the original figureof the type (Fig. 1C, D; Alcock & Anderson, 1895: pl. 14
fig. 4, 4a) as well as Alcock (1901: pl. 6 fig. 24) (Fig. 1A,B) in most respects, except that the subhepatic region has
only a distinct spine with one sharp granule posterior toit (Figs. 5E, 7G) rather than with two distinct spines (cf.
Alcock & Anderson, 1895: pl. 14 fig. 4; Guinot & Richer
de Forges, 1995: fig. 56c, d). The USNM specimen on theother hand, possesses two distinct spines on the subhepatic
region (Figs. 5F, 7H). The proportions and armature of
the P2–P4 of both specimens (Fig. 2) agree well with theoriginal figures (Fig. 1).

Material examined. 1 male (11.8 × 9.9 mm) (OUMNH
2008-09-0039), Laccadive Sea, 12°5′35′′N 71°35′50′′E,
India, 1580–1610 m, coll. 1894, from Indian Museum. –1 female (15.6 × 13.0 mm) (USNM 42696), station 192,
Arabian Sea, 15°11′N 72°28′45′′E, 1668–1703 m, coll.
RIMSS Investigator.
Remarks. Wood-Mason (in Wood-Mason & Alcock,
1891: 269) described Hypsophrys superciliosa fromINVESTIGATOR station 105 in the Arabian Sea, Indian
Ocean (off Goa, India: 15°2′N 73°34′E) from a depth of 1353
m. He observed that he had “Four specimens, two males
and two females, of which only one pair is in good order”
and he provided measurements for the intact male (16.3 ×
13.3 mm) and female (20.5 × 17.0 mm) (Wood-Mason, in
Wood-Mason & Alcock, 1891: 270). The current depositoryof the material is not known but presumably in the IndianMuseum. Guinot & Richer de Forges (1995: 445) examinedan ovigerous female from the Maldives. The present OUMNH
specimen was obtained from the area between the Laccadive
Sea and Arabian Sea, the geographical co-ordinates being
quite close to the type locality, but it is not a syntype. The
present USNM specimen is from the Arabian Sea but it isfrom a different station and as such, cannot be a syntype
as well.

The records of “Lamoha superciliosa” from the South China
Sea and Taiwan by Serène & Lohavanijaya (1973), Richer
de Forges & Ng (2008) and Ahyong et al. (2009) are allhere regarded as belonging to L. longirostris. On the basisof the good series of specimens of L. longirostris we haveexamined from various parts of the Pacific, we now knowthat the two primary characters used by Guinot & Richer
de Forges (1995) and Richer de Forges & Ng (2008) toseparate the two species – the number of subhepatic spines(one or two) and the armature of the supraorbital margin(with one or no spine) are variable (see remarks for next
species). Richer de Forges & Ng (2008) also commented
that the black spot on the cheliped is longer in L. longirostriscompared to L. superciliosa from the South China Sea andTaiwan, but we now find that this character varies as well.
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Fig. 2. Lamoha superciliosa (Wood-Mason, in Wood-Mason &Alcock, 1891). A, C–G, male (11.8 × 9.9 mm) (OUMNH 200809-0039), Laccadive Sea; B, H, female (15.6 × 13.0 mm) (USNM
42696), Arabian Sea. A, B, overall habitus; C, left G1, ventral
view; D, left G1, dorsal view; E, left G2; F, outer view of rightchela; G, H, left P4 dactylus.

Fig. 3. Lamoha longirostris (Chen, 1986). A, paralectotype male
(22.9 × 18.6 mm) (ZRC 1999.0007), East China Sea; B, male (29.2
× 24.2 mm) (ZRC 2008.990), Philippines; C, female (24.9 × 20.3mm) (ZRC 2016.0556), South China Sea.

Ocean while the latter only from the West Pacific) and thedistinctiveness of the ambulatory dactyli indicate that bothare separate species.

The only characters that appear to separate L. superciliosa s.
str. from L. longirostris is the general form of the carapace
and ambulatory legs. In L. superciliosa s. str., the carapace
is relatively more quadrate (Figs. 2A, B, 5E, F) (carapace
slightly more subovate with the lateral margins usually moreconverging in L. longirostris, Figs. 3, 4A, B, 5A–D); thegastric regions are relatively smoother (Figs. 1A, B, 5E, F)(gastric regions more uneven, with low swellings and some
low tubercles in L. longirostris, Figs. 3, 4A, B, 5A–D); andmore importantly, the P2–P4 dactylus is proportionately
longer and more slender (Fig. 2A, B, G, H) (relatively shorterand stouter in L. longirostris, Figs. 3, 4). The proportions of
the ambulatory dactylus are surprisingly constant regardlessof sex or size, being always relatively shorter and stouter in thegood series of L. longirostris examined here. The relatively
longer and more slender dactylus of L. superciliosa s. str. isalso evident even in the female figured by Guinot & Richer
de Forges (1995: fig. 56c) from the Maldives. The G1 (Fig.
2C, D) and G2 (Fig. 2E) structures of L. superciliosa arealmost identical to those figured for L. longirostris by Ng& Chen (1999: figs. 1j, k, 2a, b).

Lamoha longirostris (Chen, 1986)(Figs. 3, 4, 5A–D, 6A–G, 7A–F, 23, 24A)
Hypsophrus longirostris Chen, 1986: 227.
Hypsophrys futuna Guinot & Richer de Forges, 1995: 456, figs.
61l, 66 a, g. – Cleva et al., 2007: 251, fig. 20A.
Lamoha superciliosa – Serène & Lohavanijaya, 1973: 30, pl. 4
fig. A. – Ahyong et al., 2009: 98, figs. 66–68. – Richer deForges & Ng, 2008: 23, figs. 19A–C, 20A–E, 21A–D, 22E, F(not Hypsophrys superciliosa Wood-Mason, in Wood-Mason& Alcock, 1891)
Lamoha longirostris – Ng & Chen, 1999: 760. – Ng et al., 2001:
54. – Takeda et al., 2005: 106, fig. 1B. – Ng et al., 2008: 40.
– Richer de Forges & Ng, 2008: 20, figs. 17A–D, 18A–G,
22B–D. – Ahyong et al., 2009: 94, figs. 61–63.

Material examined. Paralectotype: male (22.9 × 18.6mm) (ZRC 1999.0007), East China Sea, St. 2V-9, 28°10′N
127°30′E, 900 m, coll. 3 January 1981. Others: 1 male (29.2
× 24.2 mm) (ZRC 2008.0990), station CP 2753, eastern
Luzon, 15°37.23′N 121°58.24′E, Philippines, 1252–1350m, trawl, coll. AURORA 2007 Expedition, 3 June 2007. –1 male (25.2 × 20.2 mm) (ZRC 1999.0410), off Tungsha
Island, Taiwan, South China Sea, trawl coll. P.-H. Ho, 23

Lamoha superciliosa s. str. is known from few specimens andwhile it is clearly close to L. longirostris, their geographicalseparation (the former is only known from the eastern Indian245

Ng & Richer de Forges: Homolidae from the South China Seamm) (USNM 1150828), station 98, Pagan Island, Northern
Mariana Islands, 18°05′48′′N 145°41′48′′E, 896 m, coll.
Townsend Cromwell, 6 May 1982. – 1 male (22.0 × 18.3mm) (USNM 1150825), station 231, Agrihan Island, Mariana
Islands, 18°47.4′N 145°35′E, 1280 m, coll. Townsend
Cromwell, 11 July 1982. – 1 ovigerous female (18.5 × 16.6mm) (MNHN-IU-2011-2732), station CP 3686, Papua New
Guinea, 03°16′S 147°18′E, 964–1025 m, coll. BIOPAPUA,
26 September 2010.
Remarks. The specimen from the East China Sea is aparalectotype (see Ng & Chen, 1999). Lamoha futuna(Guinot & Richer de Forges, 1995) was synonymised with L.
longirostris by Ng & Chen (1999) after examining the typesof both species. The distribution of this deep-sea species istherefore very wide, from the South China Sea and China
to the central Pacific.
The prominent black spot on each side of the propodus of
their chelipeds (see Richer de Forges & Ng, 2008: fig. 18C,
D) is distinct in all specimens, but varies in size. This spot
was “interpreted” by Williams (1976) as being a luminescentorgan. Nobody, however, has observed this phenomenon ona live animal.
In L. longirostris, the supraorbital margin varies fromalmost entire, without trace of any tooth or spine (Fig. 6G),
possessing a low tooth (Fig. 6A, B, E, F) or distinct spineor tooth (Fig. 6C, D). Both specimens of L. superciliosa
have a small tooth on the supraorbital margin (Fig. 6H). The
subhepatic region of L. longirostris always has one distinct
spine and there is sometimes also a sharp granule or a small
spine present on the outer part as well, but always distinctly
smaller than the inner one (Figs. 5A, 7A, C, E, F). Manyof the specimens of L. longirostris, however, have only one
spine, with the remainder of the surface smooth (Fig. 5B–D,
7B, D). In L. superciliosa, however, this second tooth iseither small (Fig. 7G) or can be as large as the inner one(Figs. 1A, C, 7H).

Fig. 4. Lamoha longirostris (Chen, 1986). A, male (15.3 × 12.6mm) (ZRC 2016.0556), South China Sea; B, C, ovigerous female(18.5 × 16.6 mm) (MNHN-IU-2011-2732), Papua New Guinea;
D, female (13.1 × 10.0 mm) (ZRC 2008.992), South China Sea.
A, B, overall habitus; C, right P4 dactylus; D, left P4 dactylus.

April 1995. – 2 ovigerous females (20.8 × 18.2 mm, 26.3
× 20.8 mm) (ZRC 2008.0991), Taiwan, South China Sea,
1400 m, coll. Taiwan University, 2000s. – 1 male (15.0 ×
12.8 mm), 1 ovigerous female (23.2 × 18.9 mm), 1 female(24.9 × 20.3 mm) (ZRC 2016.0555), station CP 4167, off
Tungsha Island, 22°06.12′N 119°07.77′E – 22°02.69′N
119°03.64′E, Taiwan, South China Sea, 1306–1756 m, mud
substrate, trawl, coll. ZHONGSHA 2015 Cruise, 1 August
2015. – 2 males (24.9 × 20.5 mm, 15.3 × 12.6 mm), 1
ovigerous female (24.2 × 20.4 mm), 1 female (23.4 × 19.4mm) (ZRC 2016.0556), station CTS 2, cold seep, 22°5.19′N
119°48.03′E – 22°2.22′N 119°48.02′E, off southern Taiwan,
South China Sea, 1360–1669 m, mud sediment with manydead bivalves and vent tubes, trawl, coll. 30 May 2015. – 1
female (17.9 × 15.0 mm) (ZRC 2008.0993), Taiwan, station
OCP 280, 24°23.71′N 122°14.22′E, 1213–1261 m, eastern
Taiwan, South China Sea, trawl, coll. TAIWAN 2005 Cruise,
14 June 2005. – 1 male (18.0 × 14.8 mm), 2 females (13.1 ×
10.0 mm, 10.8 × 8.5 mm) (ZRC 2008.0992), station CP 277,
24°23.57′N 122°14.12′E, 1222–1261 m, southern Taiwan,
South China Sea, trawl, coll. TAIWAN 2005 Cruise, 14
June 2005. – 1 male (13.8 × 11.2 mm), 2 juvenile females(11.8 × 9.1 mm, 10.0 × 7.2 mm) (ZRC 2016.0557), station
CST 17, muddy sediment, Pointer Ridge, Taiwan, 22°3.79′N
118°58.80′E – 22°3.78′N 119°4.11′E, 1482 m coll. 1 May
2016. – 2 ovigerous females (28.6 × 25.3 mm, 27.1 × 23.9

Colour. In life, the carapace, chelipeds and ambulatory legs
are red to pink, with the fingers dark-brown to almost black(Figs. 23, 24A) (see also Richer de Forges & Ng, 2008: fig.
18; Ahyong et al., 2009: figs. 61, 62, 66, 67).
Ecology. A specimen of L. longirostris was observed bya submersible at a depth of 1212 m in the Mariana Islandgroup (18.323°N 165.979°E) on 13 August 2016 carrying
an unidentified sea anemone (Fig. 23). Another Lamoha
species, L. inflata (Guinot & Richer de Forges, 1981) has
also been observed to use the P5 to cling to and manipulate
sea anemones. Chintiroglou et al. (1996) first reported
that specimens of L. inflata caught in deep-water traps
often carried a species of Isanthus (Anthozoa, Actiniaria,
Isanthidae). They noted the crabs were caught from hardbottoms and suggested that because the “special chelate
structure of the last pair of legs which is raised over the
carapace, that the crab must be the active partner, probably
manipulating the anemone with the P5 to take it off thesubstratum and to establish the association.” (Chintiroglou
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Fig. 5. Carapace. A–D, Lamoha longirostris (Chen, 1986). A, paralectotype male (22.9 × 18.6 mm) (ZRC 1999.0007), East China Sea;
B, male (29.2 × 24.2 mm) (ZRC 2008.990), Philippines; C, female (24.9 × 20.3 mm) (ZRC 2016.0555), South China Sea; D, male (15.3
× 12.6 mm) (ZRC 2016.0556), South China Sea; E, F, L. superciliosa (Wood-Mason, in Wood-Mason & Alcock, 1891). E, male (11.8 ×
9.9 mm) (OUMNH 2008-09-0039), Laccadive Sea; F, female (15.6 × 13.0 mm) (USNM 42696), Arabian Sea.

Material examined. 1 ovigerous female (24.8 × 21.8 mm)(ZRC 2009.0786), station P4, Balicasag, Panglao, Bohol,
Visayas, Philippines, 9°31.1′N 123°41.5′E, coll. 31 May
2004. – 1 male (21.8 × 18.9 mm) (ZRC 2001.2276), NE
Taiwan, Ilan Province, Nangfangao, Shu-Ao, coll. J.-F.
Huang, 3 July 1992. – 1 male (20.4 × 18.4 mm) (ZRC
2016.0200), station CP 4153, north of Macclesfield Bank,
16°13.94′N 114°27.21′E – 16°14.24′N 114°29.55′E, Taiwan,
South China Sea, 318 m, sponge substrate with sea pensand echinoderms, trawl, coll. ZHONGSHA 2015 Cruise,
27 July 2015.

et al., 1996: 22) (see also Guinot et al., 1995). The present
photographs of a L. longirostris photographed in situ (Fig.
23) in his natural hard bottom habitat carrying an unidentified
anemone confirms their hypothesis. Another homolid,
Paramola japonica (Parisi, 1915) is also reported to carry sea
anemones (see Wicksten, 1985; Guinot & Wicksten, 2015).
Lamoha murotoensis (Sakai, 1979)(Fig. 24B)
Hypsophrys murotoensis Sakai, 1979: 6. – Guinot & Richer deForges, 1981: 541. – Miyake, 1983: 197. – Guinot & Richer
de Forges, 1995: 449.
Hypsophrys longipes – Matsuzawa, 1977: pl. 87, figs. 1, 2. – Sakai,
1977: 54.
Lamoha murotoensis – Tan et al., 2000: 185. – Ng et al., 2001:
6. – Ng & Wang, 2002: 14. – Richer de Forges & Ng, 2008:
21. – Ahyong et al., 2009: 96.

Colour. In life, the carapace, chelipeds and ambulatory legs
are pink, with the stronger cristae and larger granules andspines on the carapace white (Fig. 24B) (see also Richer deForges & Ng, 2007: fig. 8B; Ahyong et al., 2009: fig. 64).
Remarks. This distinctively coloured species is widelydistributed from Japan to Madagascar (Ahyong et al., 2009).

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Fig. 6. A–G, Lamoha longirostris (Chen, 1986). A, paralectotype male (22.9 × 18.6 mm) (ZRC 1999.0007), East China Sea; B, male (24.9
× 20.5 mm) (ZRC 2016.0556), South China Sea; C, male (29.2 × 24.2 mm) (ZRC 2008.990), Philippines; D, ovigerous female (24.2 ×
20.4 mm) (ZRC 2016.0556) South China Sea; E, female (24.9 × 20.3 mm) (ZRC 2016.0555), South China Sea; F, female (23.4 × 19.4mm) (ZRC 2016.0556), South China Sea; G, male (15.3 × 12.6 mm) (ZRC 2016.0556), South China Sea. H, L. superciliosa (Wood-Mason,in Wood-Mason & Alcock, 1891), male (11.8 × 9.9 mm) (OUMNH 2008-09-0039), Laccadive Sea.

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Fig. 7. A–G, Lamoha longirostris (Chen, 1986). A, paralectotype male (22.9 × 18.6 mm) (ZRC 1999.0007), East China Sea; B, male (24.9
× 20.5 mm) (ZRC 2016.0556), South China Sea; C, male (29.2 × 24.2 mm) (ZRC 2008.990), Philippines; D, female (24.9 × 20.3 mm)(ZRC 2016.0555), South China Sea; E, female (23.4 × 19.4 mm) (ZRC 2016.0556), South China Sea; F, male (15.3 × 12.6 mm) (ZRC
2016.0556), South China Sea; G, L. superciliosa (Wood-Mason, in Wood-Mason & Alcock, 1891), male (11.8 × 9.9 mm) (OUMNH
2008-09-0039), Laccadive Sea. H, L. superciliosa (Wood-Mason, in Wood-Mason & Alcock, 1891), female (15.6 × 13.0 mm) (USNM
42696), Arabian Sea.

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Paromola Wood-Mason, in Wood-Mason & Alcock,
1891

Moloha majora (Kubo, 1936)(Fig. 24E)

Paromola macrochira Sakai, 1961
(Fig. 24C, D)

Latreillopsis major Kubo, 1936: 63, pl. 17 (see Guinot & Richer
de Forges, 1995: 384; Richer de Forges & Ng, 2007: 34; forcomplete synonymy, excluding Philippine records).

Paromola macrochira Sakai, 1961: 146, pl. 4, fig. 5. – Griffin,
1965: 87 (part); Serène & Lohavanijaya, 1973: 26, 27, fig.
28 (list). – Sakai, 1976: 40, pl. 10. – Jenkins, 1977: 4 (list). –Matsuzawa, 1977: pl. 85, figs. 1, 2. – Miyake, 1983: 13, 197,
pl. 5 fig. 2. – Guinot & Richer de Forges, 1995: 367: figs.
21c–e, 27 f. – Ng & Huang, 1997: 262, fig. 1B, C. – Ikeda,
1998: 68–69, pl. 10. – Miyake, 1998: 13, 197, pl. 5 fig. 2. –Muraoka, 1998: 15; Tan et al., 2000: 186. – Ng et al., 2001:
6. – Chen & Sun, 2002: 140, fig. 58. – Marumura & Kosaka,
2003: 21. – Richer de Forges & Ng, 2007: 36. – Ahyong etal., 2009: 111, figs. 78–81. – Ng et al., 2008: 41. – Ng, 2015:
1447, figs. 1–5.
Paromola alcocki faughni Serène & Lohavanijaya, 1973: 27, fig.
32–38, pl. 3C. – Kensley, 1980: 25.
Moloha ? alcocki faughni – Guinot & Richer de Forges, 1995:
376, 383.

Material examined. 1 male (54.8 × 47.8 mm, broken)(ZRC 2016.0197), station CP 4153, north of Macclesfield
Bank, 16°13.94′N 114°27.21′E – 16°14.25′N 114°29.55′E,
Taiwan, South China Sea, 318 m, sponge substrate withsea pens and echinoderms, trawl, coll. ZHONGSHA 2015
Cruise, 27 July 2015.
Colour. The colour is typical for the species, being orangishred overall (Fig. 24E) (see also Ahyong et al., 2009: fig. 73).
Remarks. Moloha majora was described from one male andone female from off Kominato in Japan by Kubo (1936).
The current location of this material is not known and may
be lost (see Ng & Richer de Forges, 2015).

Material examined. 1 female (soft, 26.5 × 22.7 mm) (ZRC
2016.0558), station DW 4095, 21°13.20′N 121°33.75′E –21°12.41′N 121°32.62′E, 517–573 m, Taiwan, South China
Sea, trawl, coll. TAIWAN 2013 Cruise, 19 May 2013. – 1juvenile (11.1 × 8.7 mm) (ZRC 2016.0559), station CP
4117, continental slope off Tungsha Island, 20°00.88′N
114°08.80′E – 20°01.87′N 114°09.36′E, Taiwan, South
China Sea, 333–421 m, coll. trawl, NANHAI 2014 Cruise,
11 January 2014. – 2 females (16.5 × 11.4 mm, 16.5 ×
11.5 mm) (ZRC 2016.0560), station CP 4155, northeast of
Macclesfield Bank, 16°13.60′N 115°01.61′E – 16°11.21′N
114°59.77′E, Taiwan, South China Sea, 510–526 m, trawl,coll. ZHONGSHA 2015 Cruise, 28 July 2015. – 1 soft malespecimen (damaged) (ZRC 2016.0561), station CP 4156,
northeast of Macclesfield Bank, 16°09.80′N 114°58.73′E –16°12.19′N 115°00.53′E, Taiwan, South China Sea, 503–511
m, trawl, coll. ZHONGSHA 2015 Cruise, 28 July 2015. –1 female (16.3 × 11.1 mm) (ZRC 2016.0562), station CP
4137, continental slope off Tungsha Island, 19°53.06′N
114°21.67′E – 19°53.03′N 114°24.74′E, Taiwan, South
China Sea, 524–536 m, trawl, coll. ZHONGSHA 2015
Cruise, 23 July 2015.

Homolochunia Doflein, 1904
Homolochunia gadaletae Guinot & Richer de Forges,
1995(Fig. 24F)
Homolochunia gadaletae Guinot & Richer de Forges, 1995: 434,
figs. 50e–f, 51d–f. – Ng & Richer de Forges, 2008: 7. – Ahyong
et al., 2009: 83, figs. 51–53.
Homolochunia valdiviae – Sakai, 1955: 106. – Sakai, 1976: 42.
– Miyake, 1983: 197. – Nagai, 1994: 50. (not Homolochunia
valdiviae Doflein, 1904)

Material examined. 1 ovigerous female (34.1 × 26.4 mm), 1
female (11.6 × 8.5 mm) (ZRC 2016.0198), station CP 4156,
northeast of Macclesfield Bank, 16°09.80′N 114°58.73′E –16°12.19′N 115°00.53′E, Taiwan, South China Sea, 503–511
m, trawl, coll. ZHONGSHA 2015 Cruise, 28 July 2015.
Colour. The carapace and pseudorostral spines are brown
to orange, with the rest of body dirty-white (Fig. 24F) (seealso Ahyong et al., 2009: figs. 51, 52).
Remarks. The female from the South China Sea fits wellwith the description of H. gadaletae. The pseudorostralspines are long, slender, curved downward and possess
two accessory spines distally. The subhepatic spines in thisspecies are sharp, with the upper one twice the length of thesecond one; and the anterolateral and posterolateral spines
are all strong and sharp. The ambulatory legs are very longand slender, with long dactyli; the P2–P4 meri all have
five spines on the dorsal margin; and the fingers of the P5
pseudochela are long and slender.

Colour. Adult Paramola macrochira are orangish-brownoverall (Ahyong et al., 2009: figs. 78, 79). The smaller
specimens obtained from the South China Sea are paleorangish-pink overall, with the ambulatory legs mostly
white (Fig. 24C, D).
Remarks. On the juveniles of this large species, the rostral
horns appear to be relatively longer than in the adults. Ng(2015) demonstrated that the Paromola alcocki faughni from
Serène & Lohavanijaya (1973) is a juvenile of P. macrochira.
Moloha Barnard, 1946
Remarks. Ng & Kumar (2015) recently reviewed Moloha
in describing a new species from India and noted that thecorrect date for the genus should be “Barnard, 1946”.
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Homolomannia Ihle, 1912

Remarks. The taxonomy of this widely distributed IndoWest Pacific species has been discussed at length by Guinot& Richer de Forges (1995).

Homolomania sibogae Ihle, 1912
(Fig. 24G)

Homologenus A. Milne-Edwards, in Henderson, 1888

Homolomannia sibogae Ihle, 1912: 208 (for complete synonymy,
see Guinot & Richer de Forges, 1995: 460; Richer de Forges& Ng, 2007: 41, fig. 8C, D; Richer de Forges & Ng, 2008:
11, figs. 9A–C, 10A–C; Ahyong et al., 2009: 91, figs. 58–60).

Remarks. Homologenus is a genus composed of small
species living in deep waters, often greater than 1000 m,
on muddy or sandy substrates. Eleven species are known
from the Atlantic and the Pacific Oceans: H. rostratus (A.
Milne-Edwards, 1880) (Caribbean Islands); H. braueri
Doflein, 1904 (Somalian coast); H. malayensis Ihle, 1912
(Indonesia); H. broussei Guinot & Richer de Forges, 1981
(French Polynesia); H. asper Zarenkov, in Zarenkov &Khodkina, 1983 (East Pacific Marcus-Necker seamounts);
H. donghaiensis Chen, 1986 (East China Sea); H. orientalis
Zarenkov, 1990 (East Pacific seamounts); H. levii Guinot &Richer de Forges, 1995 (New Caledonia); H. wallis Guinot& Richer de Forges, 1995 (Wallis and Futuna Islands); H.
boucheti Guinot & Richer de Forges, 1995 (Madere, eastern
Atlantic); and H. namakae Ng, 2016 (Hawaii). Two new
species from the South China Sea (H. exilis n. sp. and H.
brevipes n. sp.) allied to H. malayensis and H. donghaiensis
are described below.

Material examined. 1 male (25.3 × 19.2 mm) (ZRC
2009.0907), Tungkang fishing port, southern Taiwan, coll.
P.K.L. Ng, 7 April 2004. – 1 male (15.4 × 10.2 mm), 2 females(12.3 × 8.6 mm, 11.4 × 8.4 mm) (ZRC 2016.0199), station
CP 4135, continental shelf of Tungsha Island, 19°58.42′N
114°32.93′E – 19°58.94′N 114°37.70′E, Taiwan, South
China Sea, 211–218 m, sandy coral substrate, trawl, coll.
ZHONGSHA 2015, 23 July 2015.
Colour. The colour pattern here, a uniform brownish-red (Fig.
24G), is typical for this species, although large individuals
tend to be brown (see also Richer de Forges & Ng, 2007:
fig. 8C, D; Ahyong et al., 2009: figs. 58, 59).
Paromolopsis Wood-Mason, in Wood-Mason & Alcock,
1891

The two new species of Homologenus described here arerepresented by both sexes, and allow us to observe somesexually dimorphic features. In females, the armature on
P2–P4 (Figs. 9A, 10A, 12E–G, 16A, B, 19A–C, I–K) is
stronger and more prominent than in males (Figs. 9B, 10B,
12I–K, 16C, 19E–G, M–O). The spines on the pleonalsomites 2–5 in males (Fig. 21A, B, D, E, G, H), however,are always stronger and more prominent than in females(Figs. 11F, 15F, 17F).

Paromolopsis boasi Wood-Mason, in Wood-Mason &Alcock, 1891
(Fig. 24H)
Paromolopsis boasi Wood-Mason & Alcock, 1891: 268, fig. 5(for complete synonymy, see Guinot & Richer de Forges,
1995: 355; Richer de Forges & Ng, 2008: 32; Ahyong et al.,
2009: 115, fig. 82).

Of interest is that all specimens of H. brevipes have beencollected from deep waters just off the narrow continental
shelf of eastern and southeastern Taiwan; whilst those of
H. exilis are from off Tungsha Island, some 400 km westsouthwest of Taiwan.

Material examined. 1 male (31.4 × 26.9 mm) (ZRC
2016.0563), station CP 4118, continental shelf off Tungsha
Island, 20°00.76′N 115°00.83′E – 20°01.28′N 115°02.12′E,
Taiwan, South China Sea, 700–723 m, trawl, coll. NANHAI2014 Cruise, 12 January 2014. – 1 male (19.8 × 17.5 mm), 1
female (20.1 × 17.4 mm) (ZRC 2016.0564), station CP 4128,
off Tungsha Island, 20°44.86′N 116°08.01′E – 20°42.28′N
116°08.01′E, Taiwan, South China Sea, 420–444 m, trawl,coll. DONGSHA 2014 Cruise, 1 May 2014. – 1 male (20.5
× 18.2 mm), 1 male (20.1 × 17.7 mm) (ZRC 2016.0565),
station CP 4137, continental slope off Tungsha Island,
19°53.06′N 114°21.68′E – 19°53.03′N 114°24.74′E, Taiwan,
South China Sea, 524–536 m, trawl, coll. ZHONGSHA 2015
Cruise, 23 July 2015. – 1 female (18.8 × 17.1 mm) (ZRC
2016.0566), station CP 4155, northeast of Macclesfield Bank,
16°13.60′N 115°01.61′E – 16°11.21′N 114°59.77′E, Taiwan,
South China Sea, 510–526 m, trawl, coll. ZHONGSHA 2015
Cruise, 28 July 2015.

Homologenus malayensis Ihle, 1912
(Figs. 8–12, 18A, B, 20A, B, 21A–C, 24I, J)
Homologenus malayensis Ihle, 1912: 209. – Ihle, 1913: 70, figs. 25,
31, 36–38, pl. 2, figs. 13–15. – Balss, 1940: fig. 98. – Guinot,
1979: 232. – Guinot & Richer de Forges, 1981: 553. – Guinot& Richer de Forges, 1995: 476, figs. 65 A, B, 66 d. – Ng etal., 2008: 40.

Material examined. 2 males (carapace widths 6.2 mm,
6.1 mm; both with broken rostrums) (ZRC 2016.0194, ex
MNHN-IU-2011-2712), station CP 3762, 03°57′S 153°49′E,
995–1050 m, Papua New Guinea, coll. BIOPAPUA, 14
October 2010. – 1 ovigerous female (11.8 × 9.6 mm; rostrum
broken) (MNHN-IU-2014-18963), station CP 4449, New
Ireland, Papua New Guinea, 02°10′S 150°11′E, 623–908
m, coll. KAVIENG 2014, 1 September 2014. – 1 ovigerous
female (14.6 × 14.0 mm) (ZRC 2016.0195, ex MNHNIU-2014-18991), station CP 4442, New Ireland, Papua New
Guinea, 02°164S 150°36′E, 892–925 m, coll. KAVIENG

Colour. The colour in life is a typical orangish-brown on the
carapace and ambulatory legs (Fig. 24H) (see also Ahyong
et al., 2009: fig. 82).

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Fig. 8. Homologenus malayensis Ihle, 1912. A, B, F, G, lectotype female (15.5 × 9.8 mm, ZMA DE 102.967), station 122,
northern Sulawesi, 1°58.5′N 125°00.5′E, Indonesia, 1264–1165 m, coll. SIBOGA Expedition, Weber, 17 July 1899 (A, B, after Ihle,
1913: pl. 2 figs. 13, 14; F, G, after Guinot & Richer de Forges, 1995: fig. 65); C, type specimen, sex not stated (after Ihle, 1913: fig.
31); D, E, paralectotype male (size not specified) (after Ihle, 1913: figs. 37, 38). A, overall habitus; B, ventral view of cephalothorax;
C, left P5 pseudochela (propodus and dactylus); D, left G1; E, left G2; F, dorsal view of carapace; G, ventral view of cephalothorax.

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Fig. 9. Homologenus malayensis Ihle, 1912, overall habitus. A,
lectotype female (15.5 × 9.8 mm) (NNM ZMA DE 102.967),
station 122, northern Sulawesi, 1°58.5′N 125°00.5′E, Indonesia,
1264–1165 m, coll. SIBOGA Expedition, Weber, 17 July 1899;
B, paralectotype (11.3 × 6.1 mm) (NNM ZMA DE 102.967),
same data as lectotype. Photographs courtesy of Charles Fransen.

Fig. 10. Homologenus malayensis Ihle, 1912, overall habitus. A,
female (16.0 × 13.9 mm) (ZRC 2016.0196), Papua New Guinea; B,
male (carapace width 6.2 mm, rostrum damaged) (ZRC 2016.0194),
Papua New Guinea.

Richer de Forges (1995: 476, fig. 65) selected a lectotype
female among the material of Ihle (1912, 1913) and figured
the carapace (Figs. 8A, B, F, G, 9A). These agree very wellwith the material from Papua New Guinea (Figs. 10–12).
The figure of the G1 by Ihle (1913: fig. 37) (Fig. 8D) was
made in situ and shows the laterally flattened distal chitinous
part; it actually agrees with other material examined here
when viewed under the equivalent orientation (Fig. 12C).

2014, 31 August 2014. – 1 ovigerous female (13.8 × 12.3mm) (MNHN-IU-2014-18813), station CP 4436, New
Ireland, Papua New Guinea, 02°16′S 150°45′E, 1128–1135m, coll. KAVIENG 2014, 30 August 2014. – 1 female (11.4
× 10.6 mm) (MNHN-IU-2014-18809), station CP 4433, New
Ireland, Papua New Guinea, 02°16′S 150°48′E, 1056–1200m, coll. KAVIENG 2014, 30 August 2014. – 1 ovigerous
female (14.9 × 9.9 mm) (MNHN-IU-2014-8052), station CP
4431, New Ireland, Papua New Guinea, 02°16′S - 150°40′E,
830–894 m, coll. KAVIENG 2014, 29 August 2014. – 1
male (10.7 × 6.2 mm) (MNHN-IU-2015-1182), station CP
4434, New Ireland, Papua New Guinea, 02°19′S 150°47′E,
1066–1200 m, coll. KAVIENG 2014, 30 August 2014. – 2
ovigerous females (16.0 × 13.9 mm, 16.1 × 12.9 mm) (ZRC
2016.0196, ex MNHN-IU-2015-80), station CP 4245, 04°06′S
148°10′E, northwest of Vitu Islands, Bismarck Sea, 790–808
m, coll. MADEEP 2014, 22 April 2014.

Guinot & Richer de Forges (1995: 476) also recorded
several specimens from Indonesia and southern Philippinesand referred them to H. malayensis on the basis of theambulatory leg armature. Considering their provenance, it
seems reasonable they also belong to this species as presently
defined. The doubtful record by Nagai (1994) from Japan ishere referred to the present new species, H. exilis.
The figures of H. malayensis by Ihle (1913: pl. 2 figs. 13, 14)(Fig. 8A, B) depict a female specimen which apparently has
slightly shorter and stouter legs than the material we haveon hand from Papua New Guinea (Fig. 10). However, thisis just the consequence of how the legs are oriented whendrawn and/or photographed. When viewed obliquely, the legs,
especially the merus, appears more slender than it actuallyis, giving the impression of being more slender. Charles
Fransen (The Naturalis, Leiden) kindly photographed thetypes of H. malayensis for us (Fig. 9) and the proportionsof their ambulatory legs agree very well with the material
here figured from Papua New Guinea (Fig. 10).

Colour. The carapace is essentially dirty white to pale pinkoverall but the numerous setae trap a great deal of sediment,
giving the animals an overall light brown appearance in life
(Fig. 24I, J).
Remarks. In their revision of the genus, Guinot & Richer
de Forges (1995) examined and figured the types andclarified the identity of H. malayensis. They characterised
it by the presence of a distinct gastric spine, absence ofdistinct spines behind the lateral spine, the distal marginof P2 to P4 armed only with a single spine and unarmed
ventral margin of P4 (Figs. 8B, 12G, K) (Guinot & Richer
de Forges, 1995: 469–470). They recorded the species from
northwest of Sulawesi in Indonesia (type locality), southernJava and southern Philippines (Guinot & Richer de Forges,
1995: 476) with a doubtful record from Japan. Guinot &In the series of specimens examined (as well as what is
known about H. malayensis, cf. Guinot & Richer de Forges,
1995), there is usually only one spine on the anterolateralangle of the merus of the third maxilliped (Fig. 8G). Twoovigerous female specimens from Papua New Guinea (13.8 ×
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Fig. 11. Homologenus malayensis Ihle, 1912, female (16.0 × 13.9 mm) (ZRC 2016.0196), Papua New Guinea. A, frontal view of
cephalothorax; B, rostrum, pseudorostrum and orbit; C, left third maxilliped; D, dorsal view of right cheliped carpus; E, outer view ofleft chela; F, posterior carapace margin and male pleonal somites 1.

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Fig. 12. Homologenus malayensis Ihle, 1912. A–D, I–L, male (carapace width 6.2 mm, rostrum damaged) (ZRC 2016.0194), Papua New
Guinea; E–H, female (16.0 × 13.9 mm) (ZRC 2016.0196), Papua New Guinea. A, left third maxilliped; B, dorsal view of right cheliped
carpus; C, left G1; D, left G2; E–L, female left P2–P5 merus, respectively; I–L, male left P2–P5, respectively. Scales: C, D = 0.5 mm;
E–L = 5.0 mm.

12.3 mm, MNHN-IU-2014-18813; 16.0 × 13.9 mm, MNHNIU-2015-80), however, have two spines (Fig. 11C), so thischaracter is not always reliable.

957–988 m, Taiwan, South China Sea, sandy bottom, trawl,coll. ZHONGSHA 2015 Cruise, 22 July 2015. Paratypes:
1 male (10.2 × 7.4 mm) (ZRC 2016.0568), same data as
holotype. – 1 female (15.1 × 11.4 mm) (ZRC 2016.0569),
station CP 4133, southwest of Tungsha Island, 20°04.51′N
116°22.37′E–19°58.89′N 116°24.28′E, 999–1070 m, Taiwan,
South China Sea, sandy bottom with thin stalked sponges,
trawl, coll. ZHONGSHA 2015 Cruise, 22 July 2015. – 1 male
(11.4 × 9.6 mm) (ZRC 2016.0570), station CST 11, muddy
bottom with numerous branching corals, Formosa Ridge,
Taiwan, 22°8.83′N 119°15.68′E – 22°6.46′N 119°17.43′E,
1176–1318 m, coll. 27 April 2016.

Homologenus exilis n. sp.
(Figs. 14, 15, 18D–F, 19A–H, 20C–E, 21D–F, 22A–C)?Homologenus malayensis – Nagai, 1994: 50, pl. 1, fig. 3. (notHomologenus malayensis Ihle, 1912)

Material examined. Holotype: ovigerous female (13.5 ×
10.4 mm) (NTOU), station CP 4132, southwest of Tungsha
Island, 20°11.46′N 116°20.14′E – 20°07.26′N 116°21.51′E,
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Fig. 14. Homologenus exilis n. sp., overall habitus. A, holotype
ovigerous female (13.5 × 10.4 mm) (NTOU), South China Sea; B,
paratype male (10.2 × 7.4 mm) (ZRC 2016.0568), South China Sea.

closed; margins of merus spinose; carpus with 4 prominent
spines; dorsal margin of chela with 4 spines, ventral margin
with 4 spines (Fig. 21F). Ambulatory legs (P2–P4) very
long, slender; P5 in dorsal position, merus without spines,
reaching base of anterolateral spine when folded on carapace,dactylus long, slender, curved, sharp, touching proximal
spine of propodus, forming pseudochela (Figs. 14, 19D,
H, 20C–E); female: P2 merus with 4 or 5 spines on dorsal
margin, outer surface with 2 spines, ventral margin with 6
spinules; P3 merus with 5 long straight spines on dorsal
margin, outer surface with 1 short spine, ventral margin
with 8 spines and spinules; P4 merus with 4 or 5 spines on
dorsal margin, outer surface with 1 spinule, ventral margin
with 5 or 6 spines (Figs. 14A, 19A–C); male: P2 merus
with 3 or 4 spines on dorsal margin, outer surface with 2
spines, ventral margin with 6 spinules; P3 merus with 4
long straight spines on dorsal margin, outer surface with
2 spinules, ventral margin with 5 spines and spinules; P4
merus with 3 spines on dorsal margin, outer surface with 1
spinule, ventral margin with 3 spines (Figs. 14B, 19E–G);
P2–P4 propodus very long, dactylus very long, falciform(Fig. 14). Armature of pleonal somites as follows: female
somite 2 with 1 median spine, somite 3 with 4 low spines,
somite 4 with 4 very low spines, somite 5 with 4 very low
spines, telson, somites 1, 5 and 6 unarmed (Fig. 15F); male
somite 2 with 1 median spine, somite 3 with 5 low spines,
somite 4 with 4 low spines, somite 5 with 4 low spines,
telson, somites 1 and 6 unarmed (Fig. 21D, E). G1 relatively
stout, distal part subtruncate (Fig. 22A, B).

Fig. 13. Homologenus donghaiensis Chen, 1986, holotype male
(11.0 × 10.3 mm) (IOCAS KY8B-71), East China Sea, 900 m, onsoft mud, 3 August 1981. A, overall habitus; B, dorsal view of left
P1–P5; C, ventral view of P2–P4. Photographs courtesy of W. Jiang.

Diagnosis. Small species, covered with scattered long andshort setae (Fig. 14). Carapace longer than wide; male
carapace longitudinally ovate; anterior half of female carapace
slightly narrower than posterior half; surface of carapace
granulous; gastro-cardiac and branchio-cardiac grooves wellmarked; with long sharp median gastric spine and 2 shortepigastric spines; short spine at angle of buccal cavity; lineof prominent granules marking border of pterygostomian
region (Figs. 14, 15A, 18D–F). Rostrum very long, curved,
sharp, with 2 short accessory pseudorostral spines pointing
anteriorly (Figs. 14, 15A, B, 18D–F). Strong pseudorostralspines in supra-ocular position gently curving laterally
outwards; supraorbital margin with short spine (Figs. 14,
15A, B, 18D–F). Basal antennal spine strong (Fig. 15A).
Subhepatic spine slender (Fig. 15A). Anterolateral spine long,
sharp, pointing obliquely outwards (Figs. 14, 15A, 18D–F).
Anteroexternal angle of merus of third maxilliped with 2
sharp curved spines (Fig. 15C). Female cheliped short; chelaslightly inflated: chela with 6 small spines on ventral margin
and 3 spines on dorsal margin; fingers long, slender, curved
inwards; carpus with 3 long spines on external face; merus
with 5 strong spines on outer margin and 6 spines on inner
margin (Fig. 15D, E). Male cheliped short; chela inflated,
triangular in cross-section, fingers closely appressed when

Etymology. From the Latin “exilis” meaning “thin or
slender”, alluding to the relatively long ambulatory legs ofthe species when compared to the other new species, H.
brevipes. Used as a noun in apposition.

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Fig. 15. Homologenus exilis n. sp., holotype ovigerous female (13.5 × 10.4 mm) (NTOU), South China Sea. A, frontal view of cephalothorax;
B, rostrum, pseudorostrum and orbit; C, left third maxilliped; D, dorsal view of right cheliped carpus; E, outer view of left chela; F,
posterior carapace margin and male pleonal somites 1–3.

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Remarks. The two new species described here aresuperficially similar and resemble H. malayensis Ihle,
1912, and were referred to this taxon initially. They also
resemble H. donghaiensis Chen, 1986 (see Ng & Chen,
1999). The most notable difference is the observation thatin H. malayensis, the ventral margin of the merus of the P4
is completely unarmed (Figs. 8B, 12G, K) whereas in thetwo new species, there are two distinct spines on the ventral
margin (Fig. 19C, G, K, O). In addition, H. malayensis also
differs from H. exilis n. sp. and H. brevipes n. sp. in having
a proportionately longer male telson (Fig. 21A vs. Fig. 21D,
G), and the G1 is less prominently curved (Fig. 12C vs. Fig.
22A, D). Compared to H. donghaiensis, the two new species
have more longitudinally ovate and more pyriform carapaces(Fig. 18D–I) (carapace distinctly more rectangular in H.
donghaiensis, Figs. 13A, 18C); the carpus of the cheliped
has relatively weaker spines (Figs. 15D, 17D) (very strong
and long in H. donghaiensis, Fig. 13A, B); and the ventral
margin of the P4 merus is armed with spines (Fig. 19C, G,
K, O) (unarmed in H. donghaiensis, Fig. 13A–C).
In addition to the above differences, H. exilis differs from
H. malayensis in that the anteroexternal angle of the merusof the third maxilliped has two spines (Fig. 15C) (usuallywith only 1 spine in H. malayensis, Figs. 8G, 12A; Guinot& Richer de Forges, 1995: fig. 65B). The proportions of
their ambulatory legs are similar (cf. Figs. 9, 10, 12E–G,
I–K, 14, 19A–C, E–G).
With regards to the very long P2–P4, H. exilis superficiallyresembles H. donghaiensis Chen, 1986. However, H. exilis
can be easily distinguished by its more pyriform male
carapace (Figs. 14, 18D–F) (more rectangular with thelateral margins subparallel in H. donghaiensis, Figs. 13A,
18C; Ng & Chen, 1986: fig. 3a); the anteroexternal angleof the merus of the third maxilliped has two spines (Fig.
15C) (only one spine in H. donghaiensis, cf. Ng & Chen,
1999: fig. 3f); the spines on the margins of the carpus of
the cheliped are relatively weak (Fig. 15D) (carpal spines
prominent and strong in H. donghaiensis, Fig. 13A, B;
Ng & Chen, 1999: fig. 3b); the P2–P4 are proportionately
shorter (Figs. 14, 19A–C, E–G) (P2–P4 distinctly longer in
H. donghaiensis, Fig. 13A; Ng & Chen, 1999: fig. 3d); theP5 merus is relatively shorter (Figs. 14, 19D, H) (P5 merus
distinctly longer in H. donghaiensis, Fig. 13A; Ng & Chen,
1999: fig. 3e); and the P5 dactylus is proportionately longer,
extending beyond the subproximal propodal spine (Figs.
14, 20C–E) (P5 dactylus relatively shorter, not reaching thesubproximal propodal spine in H. donghaiensis, Fig. 13A, B;
Ng & Chen, 1999: fig. 3e). The distal part of the G1 of H.
exilis also appears to be slightly more truncate and stouter(Fig. 22A, B) compared to that of H. donghaiensis (cf. Ng& Chen, 1999: fig. 3h).

Fig. 16. Homologenus brevipes n. sp., overall habitus. A, holotype
ovigerous female (15.0 × 10.7 mm) (NTOU), Taiwan; B, paratypeovigerous female (15.4 × 13.5 mm) (ZRC 2016.0572), Taiwan; C,paratype male (7.6 × 5.9 mm) (ZRC 2016.0574), Taiwan.

(Fig. 15A) (proportionately shorter in H. brevipes, Fig. 17A);
the anteroexternal angle of the merus of the third maxilliped
has two spines (Fig. 15C) (only one spine in H. brevipes,
Fig. 17C); the distinctly longer P2–P4 (Figs. 14, 19A–C,
E–G) (relatively shorter P2–P4 in H. brevipes, Figs. 16,
19I–K, M–O); the P5 merus, when folded, reaches to thebase of the anterolateral spine (Fig. 14) (not reaching baseof anterolateral spine in H. brevipes, Fig. 16); and the P5
dactylus is relatively more slender and longer (Fig. 20C–E)(P5 dactylus relatively shorter and stouter in H. brevipes, Fig.
20F–H). The distal part of the G1 of H. exilis (Fig. 22A, B)
is also relatively more truncate and stouter than that of H.
brevipes (Fig. 22D, E). The anterolateral spines of H. exilis
are also relatively longer and directed more laterally (Fig.
18D–F) compared to those of H. brevipes which are shorterand directed more anteriorly (Fig. 18G–I). In general, in most
specimens, the carapace and P2–P4 in H. brevipes are moresetose (e.g., Figs. 16, 19I–P) compared to the condition in
H. exilis (e.g., Figs. 14, 19A–H).

Homologenus exilis different from H. brevipes by the
proportionately longer rostrum (Figs. 14, 15B, 18D, E)(distinctly shorter in H. brevipes, Figs. 16, 17B, 18G–I); thepseudorostral spines are gently curving laterally (Fig. 18D–F)(spines straight and directed obliquely laterally in H. brevipes,
Fig. 18G–I); the proportionately longer median gastric spine

The Japanese specimen from Wakayama Prefecture figured
by Nagai (1994: pl. 1 fig. 3) as “Homologenus malayensis”
appears to be closer to H. exilis on the basis of its relatively
longer rostrum and ambulatory legs.

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Fig. 17. Homologenus brevipes n. sp., holotype ovigerous female (15.0 × 10.7 mm) (NTOU), Taiwan. A, frontal view of cephalothorax;
B, rostrum, pseudorostrum and orbit; C, left third maxilliped; D, dorsal view of right cheliped carpus; E, outer view of left chela; F,
posterior carapace margin and male pleonal somites 1–3.

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Fig. 18. Carapace of Homologenus species. A, H. malayensis Ihle, 1912, male (carapace width 6.2 mm, rostrum damaged) (ZRC 2016.0194),
Papua New Guinea; B, H. malayensis Ihle, 1912, female (16.0 × 13.9 mm) (ZRC 2016.0194), Papua New Guinea; C, H. donghaiensis
Chen, 1986, holotype male (11.0 × 10.3 mm) (IOCAS KY8B-71), East China Sea (photograph: W. Jiang); D, H. exilis n. sp., paratypemale (10.2 × 7.4 mm) (ZRC 2016.0568), South China Sea; E, H. exilis n. sp., holotype ovigerous female (13.5 × 10.4 mm) (NTOU),South China Sea; F, H. exilis n. sp., paratype female (15.1 × 11.4 mm) (ZRC 2016.0569), South China Sea; G, H. brevipes n. sp., paratypemale (7.6 × 5.9 mm) (ZRC 2016.0574), Taiwan; H, H. brevipes n. sp., holotype ovigerous female (15.0 × 10.7 mm) (NTOU), Taiwan;
I, H. brevipes n. sp., paratype ovigerous female (15.4 × 13.5 mm) (ZRC 2016.0572), Taiwan.

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Fig. 19. Left P2–P5 merus of Homologenus species. A–D, H. exilis n. sp., holotype ovigerous female (13.5 × 10.4 mm) (NTOU), South
China Sea; E–H, H. exilis n. sp., paratype male (10.2 × 7.4 mm) (ZRC 2016.0568), South China Sea; I–L, H. brevipes n. sp., holotype
ovigerous female (15.0 × 10.7 mm) (NTOU), Taiwan; M–P, H. brevipes n. sp., paratype male (7.6 × 5.9 mm) (ZRC 2016.0574), Taiwan.
A–D, I–L, female left P2–P5 merus, respectively; E–H, M–P, male left P2–P5, respectively. Scales = 5.0 mm.

Homologenus brevipes n. sp.
(Figs. 16, 17, 18G–I, 19I–P, 20F–H, 21G–I, 22D–F)

mm, rostrum broken) (ZRC 2013.0298), station PCP 344,
22°15.95′N 120°0.110′E, 995–1073 m, coll. TAIWAN 2006
Cruise, 8 March 2006. – 1 ovigerous female (16.8 × 15.0 mm)(ZRC 2009.1161), station CP 141, 22°12.04′N 119°59.96′E,
985–1110 m, eastern Taiwan, trawl, TAIWAN 2001 Cruise,
24 November 2001. – 1 male (12.7 × 10.0 mm) (ZRC
2009.1162), station CD 136, 22°7.75′N 120°0.87′E, 998–1211
m, eastern Taiwan, trawl, coll. TAIWAN 2001 Cruise, 22
November 2001. – 3 ovigerous females (17.0 × 15.0 mm, 17.4
× 14.3 mm, 16.8 × 13.9 mm) (ZRC 2009.1160), station CP
134, 22°16.56′N 120°6.11′E, 736–1040 m, eastern Taiwan,

Homologenus malayensis – Ho et al., 2004: 642, fig. 1A. – Ahyong
et al., 2009: 86, fig. 54, 55. (not Homologenus malayensis
Ihle, 1912)

Material examined. Holotype: ovigerous female (15.0 ×
10.7 mm) (NTOU), station PCP 344, 22°15.95′N 120°0.11′E,
995–1073 m, southern Taiwan, trawl, coll. TAIWAN 2006
Cruise, 8 March 2006. Paratypes: 1 ovigerous female (14.8261

Ng & Richer de Forges: Homolidae from the South China Sea

Fig. 20. Right P5 pseudochela (propodus and dactylus) of Homologenus species. A, H. malayensis Ihle, 1912, female (16.0 × 13.9 mm)(ZRC 2016.0196), Papua New Guinea; B, H. malayensis Ihle, 1912, male (carapace width 6.2 mm, rostrum damaged) (ZRC 2016.0194),
Papua New Guinea; C, H. exilis n. sp., holotype ovigerous female (13.5 × 10.4 mm) (NTOU), South China Sea; D, H. exilis n. sp., paratype
female (15.1 × 11.4 mm) (ZRC 2016.0569); E, H. exilis n. sp., paratype male (10.2 × 7.4 mm) (ZRC 2016.0568), South China Sea; F, H.
brevipes n. sp., holotype ovigerous female (15.0 × 10.7 mm) (NTOU), Taiwan; G, H. brevipes n. sp., paratype ovigerous female (15.4 ×
13.5 mm) (ZRC 2016.0572), Taiwan; H, H. brevipes n. sp., paratype male (7.6 × 5.9 mm) (ZRC 2016.0574), Taiwan.

trawl, coll. TAIWAN 2001 Cruise, 22 November 2001. – 1
ovigerous female (15.4 × 13.5 mm) (ZRC 2016.0572), station
CD 192, 22°17.19′N 120°1.01′E, 960–1302 m, southern
Taiwan, trawl, coll. TAIWAN 2002 Cruise, 28 August 2002.
– 1 ovigerous female (13.8 × 10.2 mm) (ZRC 2016.0573),
station PCP 445 (NTOU B00069), 22°17.10′N 120°0.17′E,
982–999 m, southern Taiwan, trawl, coll. TAIWAN 2006
Cruise, 14 July 2008. – 1 ovigerous female (12.4 × 9.9 mm,
broken rostrum), 1 male (7.6 × 5.9 mm) (ZRC 2016.0574),
southern Taiwan, 22°17.17′N 120°00.18′E – 22°13.24′N
120°00.32′E, coll. trawl, 5 July 2013. – 1 ovigerous female(16.1 × 10.9 mm) (ZRC 2016.0575), station OCP 280,
24°23.71′N 122°14.22′E, 1213–1261 m, eastern Taiwan,
trawl, coll. TAIWAN 2005 Cruise, 14 June 2005.
Diagnosis. Small species, covered with numerous longand short setae (Fig. 16). Carapace longer than wide; male
carapace longitudinally ovate; anterior half of female carapace
distinctly less wide than posterior half; surface of carapace
granular; gastro-cardiac and branchio-cardiac grooves wellmarked; with short, sharp median gastric spine and 2 shortepigastric spines; short spine at angle of buccal cavity; line
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Fig. 21. A–C, Homologenus malayensis Ihle, 1912, male (carapace width 6.2 mm, rostrum damaged) (ZRC 2016.0194), Papua New
Guinea; D–F, H. exilis n. sp., paratype male (10.2 × 7.4 mm) (ZRC 2016.0568), South China Sea; G–I, H. brevipes n. sp., paratype male
(7.6 × 5.9 mm) (ZRC 2016.0574), Taiwan. A, D, G, telson and pleonal somites 5 and 6; B, E, H, pleonal somites 2–6; C, F, I, outerview of left chela.

of prominent granules marking border of pterygostomian
region (Figs. 16, 17A, 18G–I). Rostrum very long, curved,
sharp, with 2 short accessory pseudorostral spines pointing
anteriorly (Figs. 16, 17A, B, 18G–I). Well-developed
pseudorostral spines in supra-ocular position almost straight,
directed obliquely anteriorly; supraorbital margin withshort spine (Figs. 16, 17A, B, 18G–I). Basal antennal spine
strong (Fig. 17A). Subhepatic spine slender (Fig. 17A).
Anterolateral spine long, sharp, pointing obliquely outwards,
relatively more anteriorly in angle (Figs. 16, 17A, 18G–I).
Anteroexternal angle of merus of third maxilliped with 1
sharp curved spine (Fig. 17C). Female cheliped short; chelaslightly inflated: chela with 5 small spines on ventral margin

and 4 spines on dorsal margin; fingers long, slender, curved
inwards; carpus with 3 long spines on external face; merus
with 8 strong spines on outer margin and 6 spines on inner
margin (Fig. 17D, E). Male cheliped short; chela inflated,
triangular in cross-section, fingers closely appressed whenclosed; margins of merus spinose; carpus with 4 prominent
spines; dorsal margin of chela with 4 spines, ventral margin
with 4 spines (Fig. 21I). Ambulatory legs (P2–P4) long,
slender; P5 reduced, in dorsal position, merus unarmed, notreaching base of anterolateral spine when folded on carapace;dactylus long, relatively wider, curved, sharp, reachingproximal spine of propodus, forming pseudochela (Figs. 16,
19L, P, 20F–H); female: P2 merus with 3 spines on dorsal
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Fig. 22. Gonopods of Homologenus species. A–C, H. exilis n. sp., paratype male (10.2 × 7.4 mm) (ZRC 2016.0568), South China Sea; D–F,
H. brevipes n. sp., paratype male (7.6 × 5.9 mm) (ZRC 2016.0574), Taiwan. A, C, D, F, ventral view; B, E, dorsal view. Scales = 0.5 mm.

Fig. 23. Lamoha longirostris (Chen, 1986) carrying an unidentified sea anemone in vicinity of Mariana Island group at a depth of 1212
m on 13 August 2016. Photographs courtesy of the U.S. National Oceanographic and Atmospheric Administration Office of Ocean
Exploration and Research.

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Fig. 24. Colour in life, A–H from South China Sea; I, J from Papua New Guinea and Bismarck Sea. A, Lamoha longirostris (Chen,
1986), female (24.9 × 20.3 mm) (ZRC 2016.0555); B, Lamoha murotoensis (Sakai, 1979), male (20.8 × 18.4 mm) (ZRC 2016.0200);
C, Paromola macrochira Sakai, 1961, female (16.5 × 11.4 mm) (ZRC 2016.0560); D, Paromola macrochira Sakai, 1961, female (16.3
× 11.1 mm) (ZRC 2016.0562); E, Moloha majora (Kubo, 1936), male (54.8 × 47.8 mm) (ZRC 2016.0197); F, Homolochunia gadaletae
Guinot & Richer de Forges, 1995, ovigerous female (34.1 × 26.4 mm) (ZRC 2016.0198); G, Homolomania sibogae Ihle, 1912, male
(15.4 × 11.3 mm) (ZRC 2016.0199); H, Paromolopsis boasi Wood-Mason, in Wood-Mason & Alcock, 1891, male (20.1 × 17.7 mm)(ZRC 2016.0565); I, Homologenus malayensis Ihle, 1912, ovigerous female (16.0 × 13.9 mm) (MNHN-IU-2015-80); J, Homologenus
malayensis Ihle, 1912, ovigerous female (14.9 × 9.9 mm) (MNHN-IU-2014-8052).

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margin, outer surface with 2 or 3 spinules, ventral margin
with 3 or 4 spinules; P3 merus with 2 or 3 spines on dorsal
margin, outer surface with 1 spinule, ventral margin with
4 spines; P4 merus with 3 spines on dorsal margin, outersurface with 2 spinules, ventral margin with 2 spines (Figs.
16A, B, 19I–K); male: P2 merus with 4 or 5 spines on dorsal
margin, outer surface with 2 spinules, ventral margin with
4 or 5 spinules; P3 merus with 4 spines on dorsal margin,outer surface with 2 spinules, ventral margin with 5 or 6
spinules; P4 merus with 4 spines on dorsal margin, outersurface with 2 spinules, ventral margin with 4 spines (Figs.
16C, 19M–O); P2–P4 propodus long, dactylus long, falciform(Fig. 16). Armature of pleonites as follows: female somite
2 with 1 median spine, somite 3 with 4 low spines, somite
4 with 4 very low spines, somite 5 with 4 very low spines,
telson, somites 1, 5 and 6 unarmed (Fig. 17F); male somite
2 with 1 median spine, somite 3 with 5 low spines, somite 4
with 4 low spines, somite 5 with 4 low spines, telson, somites
1 and 6 unarmed (Fig. 21G, H). G1 relatively stout, distal
part tapering to cone-like structure (Fig. 22D, E).

not be examined but the colour figure leaves no doubt it isconspecific with H. brevipes.
ACKNOWLEDGEMENTS
The authors are grateful to Chan Tin-Yam (NTOU) for
passing the interesting material to us for this study. Rafael
Lemaitre and Karen Reed (USNM) helped arrange theloan of several important lots from the Pacific. Sammy De
Grave (OUMNH) kindly loaned us the specimen of Lamoha
superciliosa. Jiang Wei (IOCAS) helped us examine andphotograph the type of Homologenus donghaiensis. Charles
Fransen (The Naturalis, Leiden) kindly helped us check thetype specimens of H. malayensis. Thanks, are due to the U.S.
National Oceanographic and Atmospheric Administration
Office of Ocean Exploration and Research for photographsof L. longirostris taken in situ in the Marianas; and to Mary
Wicksten for first bringing them to our attention. Constructive
comments on the manuscript from Shane Ahyong and Tohru
Naruse are much appreciated. The second author’s workin Singapore was the result of a visiting scientist programof the Lee Kong Chian Natural History Museum, National
University of Singapore.

Etymology. The name alludes to the relatively shorter
ambulatory legs of this species when compared to those of
the allied new species, H. exilis. Used as a noun in apposition.

LITERATURE CITED

Colour. The specimen illustrated in colour by Ahyong et al.
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In life, the carapace is dirty white overall with the anterior
part of the carapace pink.

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Remarks. As discussed earlier, there are several differences
between H. brevipes n. sp. and H. malayensis Ihle, 1912.
The most significant is in the ventral margin of the P4 merusin H. brevipes n. sp. possessing two spines (Fig. 19K, O)(margin unarmed in H. malayensis, Figs. 8B, 12G, K). In
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Fig. 13A, B); the spines on pleonal somites 2 and 3 arelow but distinct (Fig. 17F) (very short or not visible in H.
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