THE RAFFLES BULLETIN OF ZOOLOGY 2013
THE RAFFLES BULLETIN OF ZOOLOGY 2013 61(1): 189–204
Date of Publication: 28 Feb.2013
© National University of Singapore
ON THE IDENTITY OF THE INDO-WEST PACIFIC LITTORAL XANTHID CRAB,
LEPTODIUS EXARATUS (H. MILNE EDWARDS, 1834)
(CRUSTACEA: DECAPODA: BRACHYURA: XANTHIDAE)
Sang-kyu Lee
School of Biological Sciences, Seoul National University, Seoul 151-747, Republic of Korea
Email: bio249@snu.ac.kr
Jose Christopher E. Mendoza
Department of Biological Sciences, Faculty of Science, National University of Singapore
14 Science Drive 4, 117543, Republic of Singapore
Email: jcmendoza@nus.edu.sg
Peter K. L. Ng
Department of Biological Sciences, Faculty of Science, National University of Singapore
14 Science Drive 4, 117543, Republic of Singapore
Email: peterng@nus.edu.sg
Won Kim
School of Biological Sciences, Seoul National University, Seoul 151-747, Republic of Korea
Email: wonkim@plaza.snu.ac.kr (Corresponding author)
ABSTRACT. — The xanthid crab Leptodius exaratus (H. Milne Edwards, 1834) has been widely reported
from the Indo-West Pacific, where it is a ubiquitous component of the intertidal and shallow subtidal fauna.
The taxonomy of this species, however, is complex and there are many synonyms, most of which have not
been adequately treated or discussed in the past. A close examination of a series of specimens throughout its
range shows that L. exaratus should be restricted to the western Indian Ocean. Populations from the eastern
Indian and western Pacific oceans should be referred to Leptodius affinis (De Haan, 1835), a species originally
described from Japan. While their carapaces are superficially similar, the male first gonopod structure is
a reliable character to distinguish these two species. Cancer (Xantho) lividus De Haan, 1835, Leptodius
nigromaculatus Serène, 1962, as well as some of the varieties of L. exaratus established by Stimpson (1907),
are considered to be junior subjective synonyms of L. affinis (De Haan, 1835). Where necessary, lectotypes
or neotypes are designated to stabilise the taxonomy of the two species treated herein.
KEY WORDS. — Xanthidae, Leptodius exaratus, Leptodius affinis, Leptodius nigromaculatus, Indo-West
Pacific
INTRODUCTION
1906a, 1906b). Although not often discussed at length, the
taxonomy of this species is actually quite complicated as
a number of synonyms have never been recognised and/or
treated (see Stimpson, 1907; Buitendijk, 1960; Forest &
Guinot, 1961; Serène, 1984). There is also some degree of
variation: Stimpson (1907) discussed the taxonomy of L.
exaratus (as Chlorodius exaratus) and recognised several
varieties from the western Pacific (particularly southern Japan
and China), although he commented that he observed some
overlap in characters.
The genus Leptodius A. Milne-Edwards, 1863, has 12
recognised species, all of which are found in the Indo-West
Pacific region, where they are often common and ubiquitous
components of the intertidal and shallow subtidal fauna
(Serène, 1968, 1984; Ng et al., 2008). The type species,
Chlorodius exaratus H. Milne Edwards, 1834, was described
using material collected from somewhere along the shores of
the Indian subcontinent. Subsequently, this species has been
reported from many localities around the Indian and western
Pacific oceans (e.g., Dana, 1852; A. Milne-Edwards, 1868,
1873; Kossmann, 1877; Haswell, 1882; Ortman, 1893; Nobili,
Recently, the authors had independently observed some
differences between the populations of L. exaratus in the
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Lee et al.: Taxonomy of Indo-West Pacific Leptodius exaratus
mm. The statistical tests were performed with IBM® SPSS®
Statistics package (version 20).
central and western Indian Ocean, and in the western Pacific.
Exhaustive examination of the available material shows
that L. exaratus should be restricted to the form found in
the central and the western Indian Ocean, and that the “L.
exaratus” widely reported from the western Pacific should
instead be referred to Leptodius affinis (De Haan, 1835). The
main morphological characters that can be used to distinguish
the species are the armament and relative lengths of the apical
lobes of the G1. Furthermore, it is also shown that L. affinis
has several synonyms in the literature, and the attending
nomenclatural questions are resolved herein.
TAXONOMY
Superfamily Xanthoidea MacLeay, 1838
Family Xanthidae MacLeay, 1838
Subfamily Xanthinae MacLeay, 1838
Leptodius exaratus (H. Milne Edwards, 1834),
sensu stricto
(Figs. 2, 4A–D)
MATERIAL AND METHODS
Cancer inaequalis, Audouin, 1826: 86 [Egypt]; Savigny, 1809: pl.
5 fig. 7 [Egypt] [not Cancer inaequalis Olivier, 1791]
Chlorodius exaratus H. Milne Edwards, 1834: 402; 1849: pl. 11
fig. 3 [India]
Leptodius exaratus, A. Milne-Edwards, 1868: 71 [Madagascar];
Richters, 1880: 148 [Mauritius, Seychelles]; Nobili, 1906a:
121 [Persian Gulf]; 1906b: 240 [Red Sea]; Rathbun, 1911: 215
[Saya de Malha Bank, Madagascar]; Lenz, 1912: 3 [Africa];
Klunzinger, 1913: 209, pl. 3, fig. 6, pl. 5, fig. 16 [Red Sea];
Bouvier, 1915: 284 [Mauritius]; Balss, 1924: 10 [Red Sea];
Pesta, 1928: 72 [Sudan]; Maccagno, 1936: 174 [Red Sea];
Ramadan, 1936: 32 [Red Sea]; Chopra & Das, 1937: 398 (in part)
[Arabian Sea, Persian Gulf]; Forest & Guinot, 1961: 63, fig. 54
[Aldabra Is.]; Guinot, 1964: 11 [Aldabra Is., Madagascar]; 1967:
265 [Indian Ocean]; Serène, 1968: 75 (in part) [Indian Ocean];
Khan, 1977: 181, pl. 1D [Pakistan]; Kensley, 1981: 44 [South
Africa]; Serène, 1984: 184, fig. 106, pl. 26 fig. A [Madagascar,
Aldabra]; Tirmizi & Ghani, 1996: 48, fig. 18 [Pakistan]; Guinot
& Cleva, 2009: 106, with figures [Egyptian Red Sea]
Actaeodes lividus Paul’son, 1875: 35, pl. 5 fig. 2 [Red Sea]
Chlorodius (Leptodius) exaratus, Kossmann, 1877: 32, pl. 2, figs.
1–6 [Red Sea]
Xantho exaratus var. typica Ortmann, 1893: 445 (in part) [Red
Sea]
Xantho (Leptodius) exaratus, Alcock, 1898: 118 [353] (in part)
[western India, Pakistan, Persian Gulf]; Stephensen, 1946: 149,
fig. 37c [Persian Gulf]; Guinot, 1958: 92 [Mayotte Is.]; Michel,
1964: 32 [Mauritius]
The material examined in this study is deposited in the
Natural History Museum and Institute, Chiba (CBM); the
Marine Arthropod Depository Bank of Korea, Seoul National
University (MADBK); the Muséum national d’Histoire
naturelle, Paris (MNHN); the Natural History Museum,
London (NHM); the National Museum of Nature and Science,
Tokyo (NMST); and the Zoological Reference Collection,
Raffles Museum of Biodiversity Research, National University
of Singapore (ZRC). Measurements are expressed as carapace
width by carapace length, in millimetres. The abbreviation
“CL”, “CW”, “coll.”, and “G1” refer to the median carapace
length, the maximum carapace width, the collector, and the
male first gonopods, respectively. The terminology used for
carapace regions follows that of Dana (1852) and Serène
(1984). For brevity, the International Code for Zoological
Nomenclature (ICZN, 1999) is referred to as the “Code”.
Drawings were made with the aid of camera lucida attached
to a Nikon SMZ800 stereomicroscope. Images were recorded
using digital SLR camera (Nikon D200, D7000), and were
developed with software (Model Helicon Focus®). For the
morphometric analysis, the frontal width, maximum width
of sternite 4, combined length of sternites 3 and 4, total
length of the G1, and length of the G1 apical lobe were
measured (see Fig. 1) using either metric dial calipers or
ocular micrometres, where applicable, to the nearest 0.1
Fig. 1. Schematic diagram showing dimensions measured: A, carapace, dorsal view; B, male thoracic sternum, ventral view; C, left G1,
external view.
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margin with wide V-shaped notch medially; ischium
subrectangular with submedian sulcus, smooth to punctate;
exopod stout, length about 4 times width.
Xantho hydrophilus, Laurie, 1915: 444, pl. 43, fig. 1 [Sudan] [not
Cancer hydrophilus Herbst, 1790]
Xantho exaratus, Monod, 1938: 125, fig. 17B [Egyptian Red Sea];
Vatova, 1943: 19 [Somalia]; Buitendijk, 1960: 331 (in part)
[South Africa]
Xantho (Leptodius) hydrophilus, Barnard, 1950: 223, fig. 41c, 42c–e
[South Africa] [not Cancer hydrophilus Herbst, 1790]
Thoracic sternum (Fig. 2B) finely granular, glabrous. Sternites
1, 2 completely fused, separated from sternite 3 by distinct
suture; sternites 3, 4 almost completely fused except for short
notches laterally, sternite 3 distinguishable from sternite 4
by shallow groove; sternite 4 large, inflated; sternites 5–8
distinct, separate, sternite 8 not visible externally. Median
longitudinal line visible externally only on central portion
of sternite 4; within sternoabdominal cavity, visible only at
posterior portion of sternite 4, complete at level of sternites
6, 7, 8. Sternal press-button situated on sternite 5, equidistant
from sutures 4/5, 5/6.
Type locality. — Coasts of India (H. Milne Edwards, 1834: 402).
Material examined. — Neotype (here designated): male (23.7 ×
16.1 mm; NHM 1881.10), Karachi, Pakistan.
Others: Pakistan – 1 ovig. female (29.4 × 19.1 mm; NHM 1881.10),
Karachi; 1 male (30.0 × 19.4 mm), 1 female (23.4 × 14.9 mm) (ZRC
2010.0073), Buleji, 24 Feb.1982. Western India – 3 males (23.3
× 13.3 mm – 35.7 × 23.2 mm), 1 female (24.1 × 15.5 mm; NHM
1889.6.17.112–115), Bombay (?), no further data. Persian Gulf – 1
female (26.6 × 17.3 mm; NHM 1979.272), Fairlakka Is., Kuwait,
coll. D.A. Clayton, 5 May 1979; 2 males (26.3 × 17.8 mm, 29.5 ×
19.0 mm; NHM 1985.55), Bandar-e-Abbas, Iran, coll. H. Fakow,
22 Feb.1976; 2 males (20.6 × 13.9 mm; 25.8 × 17.1 mm, with
sacculinid), 1 female (20.3 ×13.7 mm; NHM 2012.1027–1029), Ras
Al Jlay’ah, Kuwait, coll. D. Clayton, 25 Oct.1979; 2 males (25.4 ×
16.7 mm, with sacculinid; 25.5 × 16.4 mm), 1 female (17.1 × 11.8
mm; NHM 2012.1030–1032), Al-Wusail, Qatar, coll. 25 Mar.1983,
don. G. Bradley; 9 males (12.2 × 8.0 mm – 29.9 × 18.9 mm), 7
females (15.5 × 11.1 mm – 20.4 × 13.2 mm; ZRC 2012.0111),
Qushm Is., Iran, coll. M. Asgari, 19 Nov.2008. Madagascar – 2
males (13.6 × 9.1 mm, 21.5 × 14.2 mm; MNHN-B6640), Nosy Be,
coll. P. Opic, 20 May 1923; 4 females (10.0 × 6.8 mm – 14.0 ×
9.0 mm; MNHN-B15992), Nosy Be, coll. A. Crosnier, Sep.1958.
Seychelles – 2 males (17.6 × 12.0 mm, 18.6 × 12.5 mm), 2 females
(17.4 × 11.6 mm, 17.6 × 11.2 mm) (MNHN-B8623), Aldabra, coll.
Calypso Expedition, May 1954.
Chelipeds (Fig. 2A) unequal. Merus with long setae on
anterior and posterior borders. Carpus finely granular, rugose
on external surface, with blunt tooth on inner angle. Palm
Description. — Carapace (Fig. 2A) transversely subovate,
about 1.5–1.6 times as broad as long; dorsal surface somewhat
convex, finely granular, rugose particularly at anterior and
lateral regions; regions well defined, separated by distinct
grooves; 2F separated from 1M by shallow, transverse groove;
2M partly divided longitudinally, 1M fused to inner branch of
2M; 3M, 2L, 3L, 4L, 5L, 6L distinct, entire; 4M indistinct;
1L very small; 1R, 2R fused, separated from 3R by indistinct
oblique groove; 1P, 2P indistinct. Front about 0.2–0.3 times as
broad as carapace breadth, not much protruded; margin deeply
sinuous medially, almost quadridentate; separated from inner
orbital tooth by notch. Orbit small, oval; superior margin with
2 fissures; inferior margin bearing 2 blunt teeth on either
side; exorbital angle separated from first anterolateral tooth
on anterolateral margin by shallow concavity. Anterolateral
margin with 4 broad, triangular teeth behind exorbital angle:
first small, acute; second broad, large; third similar to second,
slightly more produced; last small, most acute. Posterolateral
margin somewhat concave. Posterior margin granular, central
region straight. Pterygostomian region granular, setose.
Antennules (Fig. 2B, C) folding transversely, slightly
obliquely. Basal article of antenna sub-rhomboidal, short,
broad; antennal flagellum entering orbital hiatus. Epistome
narrow; central region with median projection, separated
from lateral regions by distinct notches. Third maxilliped
completely covering buccal orifice; merus subquadrate,
granular, anterolateral angle slightly produced, anterior
Fig. 2. Leptodius exaratus (H. Milne Edwards, 1834), neotype,
male, 23.7 × 16.1 mm (NHM 1881.10), Karachi, Pakistan. A, dorsal
view; B, ventral view; C, anterior view.
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inflated, rugose dorsally, smooth ventrally. Fingers stout,
darkly pigmented throughout length, except at tips with
white colour, pigment on fixed finger extending minimally
into palm; gape moderately wide; cutting margins irregularly
dentate; finger tips spoon-shaped, hollowed out, with tufts
of setae.
Mauritius (Richters, 1880; Bouvier, 1915; Michel, 1964);
Seychelles (Richters, 1880; Bouvier, 1915; Forest & Guinot,
1961; Guinot, 1964; Michel, 1964; Serène, 1984); Pakistan
(Alcock, 1898; Khan, 1977; Tirmizi & Ghani, 1996); Saya
de Malha (Rathbun, 1911); Africa (Lenz, 1912); Somalia
(Vatova, 1943); South Africa (Barnard, 1950; Buitendijk,
1960; Kensley, 1981); Mayotte (Guinot, 1958)
Ambulatory legs (Fig. 2A) short, stout; anterior margins
finely granular; anterior and posterior margins of meri
with long setae; carpi, propodi subequal in length, sparsely
setose; dactyli tomentose, spinose, ending distally in long,
chitinous claw.
Persian Gulf: Persian Gulf (Alcock, 1898; Nobili, 1906a;
Chopra & Das, 1937; Stephensen, 1946)
Red Sea: Egypt (Savigny, 1809; Audouin, 1826; Monod,
1938; Guinot & Cleva, 2009); Red Sea (Paul’son, 1875;
Kossmann, 1877; Ortmann, 1893; Nobili, 1906b; Klunzinger,
1913; Balss, 1924; Maccagno, 1936; Ramadan, 1936); Sudan
(Laurie, 1915; Pesta, 1928)
Male abdomen (Fig. 2B) narrow; somites 3–5 fused, sutures
vaguely discernible; somite 6 long, median length about 1.6
times that of telson, distal half slightly broader than proximal
half, lateral margins slightly concave. Telson subtriangular,
tip broadly rounded; tip not reaching level of sternal condyles
of P1 coxae.
Leptodius affinis (De Haan, 1835)
(Figs. 3, 4E–I)
G1 (Figs. 4A–D) long, slender, with 6–8 stout, curved
subdistal spines; with elongate apical lobe set at slight
angle with rest of G1, with 8-10 mushroom-shaped marginal
outgrowths; apical lobe about 0.05–0.07 times total length.
G2 about quarter length of G1.
Cancer (Xantho) affinis De Haan, 1835: 48, pl. 13 fig. 8 [Japan];
Krauss, 1843: 30 [Japan]
Cancer (Xantho) lividus De Haan, 1835: 48, pl. 13 fig. 6 [Japan]
[not Cancer lividus Latreille, in Milbert, 1812].
Chlorodius exaratus, Dana, 1852: 208 [Pacific] [not Chlorodius
exaratus H. Milne Edwards, 1834]
Leptodius exaratus, A. Milne-Edwards, 1873: 222 [New Caledonia];
Miers, 1879: 31 [Korea and Japan]; Haswell, 1882: 60
[Australia]; De Man, 1887a: 33 [Mergui Archipelago]; 1887b:
285 [Nordwachter Island, Java Sea]; 1892: 278 [Sulawesi];
Alcock & Anderson, 1894: 200 [Bay of Bengal, Laccadive Sea];
Balss, 1922: 127 [Japan]; Shen, 1932: figs. 57, 58c, d; 1937:
307 (list) [northern China]; Yokoya, 1933: 189 [Japan]; Sakai,
1934: 309; 1936: 151, pl. 45, fig. 3; 1965: 140, pl. 70, fig. 6;
1976: 423, pl. 153, fig. 1 [Japan]; Chopra & Das, 1937: 398
(in part) [Bay of Bengal, Mergui Archipelago]; Sankarankutty,
1962: 129 [Andaman Is.]; 1966: 351 [Sri Lanka]; Kim, 1970:
14 [Korea]; 1973: 380, fig. 144, pl. 82, fig. 109 [Korea]; Takeda
& Nunomura, 1976: 70 [New Caledonia]; Takeda, 1976: 74
[Palau]; 1978: 39 [Amakusa, Japan]; Takeda & Miyake, 1976:
109 [Ogasawara, Japan]; Yamaguchi et al., 1976: 37 [Amakusa,
Japan]; Garth & Kim, 1983: 570 [Philippines]; Dai et al., 1986:
271, figs. 154(3), 155A(1), pl. 37(4), 37(5) [China]; Dai & Yang,
1991: 292, pl. 37(4), fig. 154(3) [China]; Yamaguchi & Baba,
1993: 446, figs. 164A, B [Japan]; Jones & Morgan, 1994: 166,
167, with figure [Australia]; Minemizu, 2000: 260, with figure
[Japan]; Davie, 2002: 550–551 [Australia]; Ng & Davie, 2002:
374 [Thailand]; Paulay et al., 2003: 504 [Marianas]; Poore,
2004: 472, fig. 150b [Australia]; Davie, 2011: 233, with figure
[Australia][not Chlorodius exaratus H. Milne Edwards, 1834]
Xantho exaratus var. typica Ortmann, 1893: 445 (in part) [Samoa;
Japan; Fiji; Caroline Is.; Australia] [not Chlorodius exaratus
H. Milne Edwards, 1834]
Chlorodius exaratus var. pictus Stimpson, 1907: 54, fig. 6 [Simoda,
Japan] [not Chlorodius exaratus H. Milne Edwards, 1834]
Chlorodius exaratus var. typicus Stimpson, 1907: 55 [China; Japan]
[not Chlorodius exaratus H. Milne Edwards, 1834].
Xantho (Leptodius) exaratus, Alcock, 1898: 118 [353] (in part)
[Mergui, Andamans, Sri Lanka, Burma, Malaysia]; Laurie, 1906:
402 [Sri Lanka]; Gravely, 1927: 146 [Gulf of Mannar]; Gordon,
1931:528, 543, fig. 22b; 1934: fig. 16b [China]; Boone, 1934:
110, pl. 58 [Australia, French Polynesia]; Balss, 1935: 133 [SW
Australia]; 1938: 41 [Nauru, Marshall Is.]; Estampador, 1937:
Remarks. — Henri Milne Edwards (1834: 402) described
Chlorodius exaratus from an unspecified number of specimens
collected from “les côtes de l’Inde”. He later provided an
illustration of this species (H. Milne Edwards, 1849: pl. 11
fig. 3), presumably of the type or from the type series. The
genus Leptodius was later established by A. Milne-Edwards
(1863) for the species.
The present specimens agree with the description and
illustrations of Leptodius exaratus (e.g., H. Milne Edwards,
1849: pl. 11 fig. 3; Barnard, 1950: 223, figs. 41c, 42c, e;
Serène, 1984: 180, pl. 26 fig. A), particularly in the form of
the G1 (e.g., Stephensen, 1946: fig. 37C, Barnard, 1950: fig.
42d; Forest & Guinot, 1961: fig. 54; Serène, 1984: fig. 106).
The type, thus far, has not been found in the MNHN, where H.
Milne Edwards’ types are deposited, despite repeated attempts
to locate it. It is reasonable to assume that the type is lost
and, therefore, a neotype must now be selected to stabilise
the complex taxonomy of this species (see Discussion). A
topotypic specimen (a male, 23.7 × 16.1 mm; NHM 1881.10)
from the western coast of the Indian subcontinent (Karachi,
Pakistan) is hereby selected as the neotype for Chlorodius
exaratus H. Milne Edwards, 1834. Differences between L.
exaratus s. str. and its close sibling L. affinis (De Haan, 1835)
are discussed under the Remarks for the latter.
Distribution. — Leptodius exaratus s. str. is found in the
Western Indian Ocean, ranging from the eastern and southern
coasts of Africa, including Madagascar, to the western coast
of India; and also in the Red Sea and the Persian Gulf. It has
been reported from the following localities:
Western Indian Ocean: Western India (H. Milne Edwards,
1834; Alcock, 1898); Madagascar (A. Milne-Edwards, 1868);
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ZRC 2000.1197), Pulau Seringat, coll. C. M. Yang & S. L. Goh,
22 Jul.1997. Taiwan – 1 male (29.3 × 18.5 mm; ZRC 1995.620),
Shihmen, Taipei, coll. C. H. Wang, 24 May 1987; 1 male (29.2 ×
18.2 mm; ZRC 1999.0590), 5 males (18.9 × 12.1 mm – 26.3 × 16.5
mm), 2 females (18.8 × 12.1 mm, 22.1 × 14.1 mm; ZRC 1999.0591),
Keelung, Magang, coll. H. H. Tan, 3 Aug.1996; 2 males (29.0 ×
18.5 mm, 23.0 × 15.1 mm; ZRC 1999.0581), He Ping Tao, coll.
Keelung, H. H. Tan, 4 Aug.1996. Thailand – 9 males (14.8 × 9.7
mm – 24.5 × 15.5 mm), 4 females (13.0 × 8.4 mm – 16.7 × 11.2
mm; ZRC 2000.1034), Cape Panwa, Phuket, coll. H. H. Tan, 17
Jan.2000; 1 male (19.3 × 12.5 mm; ZRC 2001.1076), Phuket, coll.
P. K. L. Ng, 17 Feb.2001. Vanuatu – 1 male (17.9 × 11.8 mm; ZRC
2012.1233), stn. H6, rocky intertidal area near wharf of Vanuatu
Maritime College, Luganville, Espiritu Santo, coll. P. Clark, 17
Sep.2006. Vietnam – 2 males (12.7 × 8.4 mm, 22.4 × 14.0 mm),
2 females (18.7 × 12.3 mm, 28.7 × 17.9 mm; ZRC 2012.0118),
Con Dau Is., coll. H. H. Tan et al., 12 Apr.2010; 8 males (9.8 ×
6.3 mm – 13.9 × 9.0 mm), 13 females (6.4 × 4.2 mm – 12.8 × 8.6
mm; ZRC 2012.0119), Con Dau Is., 14 Apr.2010.
525; 1959: 79 [Philippines]; Sakai, 1939: 464, pl. 58, fig. 3,
pl. 91 [Japan]; Miyake, 1939: 209; 1940: 155 [Micronesia];
Chang, 1963: 99 [Taiwan]; McNeill, 1968: 58 [Australia] [not
Chlorodius exaratus H. Milne Edwards, 1834]
Xantho exaratus, Holthuis, 1953: 27 [Gilbert Is.; Tuamotu
Archipelago]; Buitendijk, 1960: 331, fig. 9 k–m (in part)
[Indonesia; China; Fiji; Japan; Myanmar; Philippines; Samoa;
Society Is.; Thailand] [not Chlorodius exaratus H. Milne
Edwards, 1834]
Leptodius nigromaculatus Serène, 1962: 255, figs. 1A–H; 1984:
182 (key) [Vietnam]; Dai et al., 1986: 272, fig. 155A [China];
Dai & Yang, 1991: 293, pl. 37(5), fig. 155A [China]; Yeh et
al., 2006: 70, figs. 1C, F, 2C–D [Taiwan]
Type locality. — Japan (De Haan, 1835: 48).
Material examined. — Australia – 2 males (21.1 × 13.8 mm;
ZRC 2012.0112), (23.4 × 15.1 mm; ZRC 2012.0113), Ningaloo
Reef, Western Australia, R. Lasley coll., 19 May 2010. China – 1
male (32.5 × 20.0 mm; ZRC 2012.0114), 3 males (13.0 × 8.5 mm
– 23.5 × 15.0 mm; ZRC 1998.542), Changpo, Hong Kong, coll. P.
K. L. Ng & S. Y. Lee, 6 Jun.1998; 2 males (30.2 × 18.6 mm, 30.7
× 19.0 mm; ZRC 1999.0625), 1 female (22.8 × 14.1 mm; ZRC
1999.0458), Nanao Is., Guangdong, coll. Y. Cai & N. K. Ng, 13
Nov.1998; 2 males (15.4 × 10.0 mm, 19.6 × 13.1 mm), 1 female
(14.2 ×9.3 mm; ZRC 2012.0115), Hainan Is., coll. Y. Cai & N. K.
Ng, 1 Dec. 1998; 2 males (17.0 × 11.6 mm, 13.2 × 8.1 mm), 2
females, (23.1 × 13.5 mm, 18.0 × 11.8 mm; ZRC 2010.0352), Shi
Jing Village, Xiamen County, Fujian Province, coll. Z. Jaafar & N.
K. Ng, 17 Nov.2005. Eastern India – 1 male (23.9 × 15.4 mm;
ZRC 2012.0110), Tranquebar, Tamil Nadu, coll. N. K. Ng, B. Y.
Lee & R. M. Lasley, Nov.2011. Indonesia – 1 female (15.8 × 10.3
mm; (ZRC 1999.1203), Bintan Is., coll. Riau, J. B. Sigurdsson, 27
Mar.1993; 1 female (13.9 × 9.5 mm; ZRC 2003.0548), Anambas
Is., stn EA-2jc7, 15 Mar.2002. Japan – 1 male (25.9 × 16.6 mm),
1 female (16.1 × 10. 6 mm; NMST-Cr 6425), Shibasaki Hayama,
Kanagawa, coll. M. Takeda, 25 Jul.1980; 2 males (23.7 × 15.2
mm, 29.4 × 19.0 mm), 1 female (21.2 × 13.5 mm; CBM-ZC 143),
2 males (12.9 × 8.6 mm, 35.0 × 21.7 mm), 1 female (20.3 × 13.3
mm; CBM-ZC 559), Ogasawara Is.; 1 female (22.6 × 14.2 mm;
ZRC 2009.0145), Iriomote Is., Yaeyama Group, southern Ryukyu
Islands, coll. N. K. Ng, 16 Jun.2000; 2 males (14.6 × 9.2 mm,
23.3 × 15.0 mm; ZRC 2011.0170), Amakusa, Kyushu, J. C. Y. Lai
coll., 3 Apr.2011. Korea – 2 males (32.0 × 20.3 mm, 36.8 × 23.2
mm; MADBK 173012_011), Iho-ri, Jejudo Is., coll. H. S. Kim, 11
Aug.1969; 3 male (19.7 × 12.7 mm – 30.6 × 19.3 mm; MADBK
173012_012), Sindo-ri, Jejudo Is., coll. S. K. Lee 25 Oct.2005; 1
male (15.7 × 10.4 mm; MADBK 173012_013), Aewel-eup, Jejudo
Is., coll. S. K. Lee, 16 Oct.2006; 2 males (20.5 × 13.2 mm, 23.1 ×
14.7 mm; MADBK 173012_014), Jo-1ri, Udo Is., Jejudo, coll. S.
K. Lee, 14 Dec.2006. Malaysia – 1 male (19.4 × 12.6 mm; ZRC
1991.461), Pulau Tioman, coll. P. K. L. Ng, 30 Mar.1982; 1 male,
(26.4 × 16.0 mm), 3 females (13.6 × 8.9 mm – 16.8 × 11.3 mm; ZRC
1985.1722-1725), Tanjong Bidara, Malacca, coll. P. K. L. Ng, 16
Feb.1985. New Caledonia – 2 males (15.8 × 10.5 mm, 17.0 × 11.4
mm), 2 females (18.3 × 12.8 mm, 17.4 × 11.4 mm; MNHN-B8631),
coll. Balansa, no date. Philippines – 2 males (23.5 × 15.6 mm, 27.7
× 18.0 mm), 1 female (21.8 × 14.1 mm; ZRC 2012.0116), 8 males
(13.6 × 9.0 mm – 29.5 × 19.3 mm), 2 females (12.6 × 8.4 mm, 23.8 ×
15.5 mm; with sacculinid; ZRC 2012.0117), Punta Taytay, Bacolod,
Negros Is., coll. J. C. E. Mendoza, 27–28 Dec.2011. Singapore
– 7 males (12.0 × 7.6 mm – 31.0 × 20.2 mm; ZRC 2012.1232),
Changi Beach Park, Singapore, 4 May 2012; 2 males (24.7 × 15.0
mm, 27.1 × 17.0 mm), 2 females (21.0 × 13.5 mm, 22.8 × 14.5
mm; ZRC 1993.33-51), Labrador Beach, Jan.1987; 3 males (19.0
× 12.1 mm – 22.7 × 14.0 mm; ZRC 1995.339), Semakau, coll. P.
K. L. Ng, 8 Feb.1993; 3 males (23.6 × 15.0 mm – 29.0 × 18.3 mm;
Description. — Carapace (Fig. 3A) transversely subovate,
about 1.4–1.6 times as broad as long; dorsal surface
depressed, finely granular, anterior, lateral regions varying
from distinctly to faintly rugose; regions well defined,
Fig. 3. Leptodius affinis (De Haan, 1835), male, 25.9 × 16.6 mm
(NSMT-Cr6425), Shibasaki, Kanagawa, Japan. A, dorsal view; B,
ventral view; C, anterior view.
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Lee et al.: Taxonomy of Indo-West Pacific Leptodius exaratus
separated by narrow, shallow grooves; 2F separated by groove
from 1M; 2M partly divided longitudinally, 1M fused to inner
branch of 2M; 3M, 2L, 3L, 4L, 5L, 6L distinct, entire; 4M
indistinct; 1L very small; 1R, 2R fused, separated from 3R
by indistinct oblique groove; 1P, 2P indistinct. Front about
0.3 times as broad as carapace breadth, not much protruded,
cut into 2 lobes, each one slightly concave near outer
side, separated from internal orbital tooth by notch. Orbit
smaller, transverse oval; superior margin with 2 fissures;
inferior margin bearing 2 blunt teeth on either side; exorbital
angle separated from first anterolateral tooth by concavity.
Anterolateral margin with 4 lobes behind exorbital angle:
first small, almost indistinguishable, depressed; second broad,
large; third similar, more prominent than second; last smallest
but most produced; separation between teeth indicated by
small sinus. Posterolateral margin somewhat concave, with
pubescence. Posterior margin granular, central region straight.
Pterygostomian region granular, setose.
subtriangular, tip broadly rounded; tip not reaching level of
sternal condyles of P1 coxae.
G1 (Figs. 4E–I) slender, long, with 5 or 6 stout, curved
subdistal spines; elongated apical lobe bordered on ventral
margin by 5 or 6 mushroom-shaped outgrowths proximally
and in distal half with 6–8 tongue-shaped outgrowths,
simple, pointed, diminishing gradually near tip. Length of
apical lobe measured from tip to subdistal region 0.11–0.14
times to total length; angle formed between apical lobe and
rest of structure relatively G1 more bent. G2 about quarter
length of G1.
Remarks. — De Haan (1835) described two species, Cancer
(Xantho) affinis and C. (X.) lividus, from Japan. These two
names were both simultaneously synonymised under Xantho
exaratus (H. Milne Edwards, 1834) var. typica by Ortmann
(1893: 445) (see also Yamaguchi & Baba, 1993). The type
material for these two names is still extant: the lectotype
(RMNH D 44644) and paralectotypes (RMNH D 42334,
RMNH D 42335, RMNH D 44646) of Cancer (Xantho) affinis
De Haan, 1835, as well as the lectotype (RMNH D 42333)
and paralectotype (RMNH D 42336) of Cancer (Xantho)
lividus De Haan, 1835, are all deposited at the Nationaal
Natuurhistorisch Museum at Leiden, The Netherlands,
and have been well documented by Yamaguchi & Baba
(1993: 446, fig. 164A, B) and Fransen et al. (1997: 116).
Stimpson (1907) noted the morphological variations among
different populations of L. exaratus in the Pacific region,
and established varieties within this species to distinguish
these populations. Serène (1962) described another species,
Leptodius nigromaculatus, from southern Vietnam, which he
thought to be more closely allied to L. gracilis (Dana, 1852),
perhaps due to the less rugose carapace and less projecting
anterolateral teeth. The holotype of L. nigromaculatus could
not be located; it has not been found in the MNHN, despite
several attempts, and it may still be in Serène’s former
institution in Nhatrang, Vietnam. Several topotypic specimens
from southern Vietnam were examined instead in order to
confirm the synonymy. Besides these, the literature is rife with
several records of “Leptodius exaratus”, or variants thereof,
from the western Pacific and eastern Indian Ocean.
Antennules (Fig. 3C) lying transversely, slightly obliquely.
Basal article of antenna sub-rhomboidal, short, broad.
Antennular flagellum occupied orbit hiatus. Epistome
narrow; central region with median projection, separated
from lateral regions by distinct notches. Third maxilliped
completely filled buccal orifice; merus subquadrate, granular,
anterolateral angle slightly produced, anterior margin with
wide V-shape notch medially; ischium subrectangular with
submedian sulcus, smooth to punctuate; exopod stout, length
about 4 times width.
Thoracic sternum (Fig. 3B) finely granular, glabrous. Sternites
1, 2 completely fused, separated from sternite 3 by distinct
suture; sternites 3, 4 almost completely fused except for short
notches laterally, sternite 3 distinguishable from sternite 4 by
shallow groove; sternite 4 large, slightly convex; sternites
5–8 distinct, separate, within not visible externally. Median
longitudinal line visible externally only on central portion
of sternite 4, complete at level of sternites 6, 7, 8. Sternal
press-button situated on sternite 5, equidistant from sutures
4/5, 5/6.
Chelipeds (Fig. 3A) asymmetrical, with a granular coating
single ended, more or less curved and pointed. Merus with
long setae on anterior and posterior borders, covered with
fine corrugation on dorsal surface. Carpus covered with
microscopic granules and fine corrugation on outer surface;
inner-distal angle bluntly round. Fingers black coloured,
with somewhat gaping between them closed, with distinctly
toothed and contiguous within; inner margins provided with
obtuse teeth, tips spoon-shaped with bristles.
Based on differences in G1 morphology, however, it is
clear that these records are not of L. exaratus s. str. Both L.
exaratus s. str. and the similar L. affinis (De Haan, 1835)
have a broad, transversely subovate carapace, which has
four broadly triangular anterolateral teeth, a rugose dorsal
surface and well-defined regions. There is much variability
and overlap in the carapace morphology within each species.
Features of the mouthparts, thoracic sternum, abdomen and
pereopods are also not useful, and morphometric analyses
of the carapace and thoracic sternum (Fig. 5A, B) reveal no
significant difference between the two species.
Ambulatory legs (Fig. 3A) smooth; meri of first to third
with setae on anterior and posterior margin; carpi, propodi
subequal in length, sparsely setose; dactyli densely covered
with short hairs, chitinous claw.
The main difference between L. exaratus s. str. and L. affinis
can be seen in the G1 morphology (Figs. 4, 5C), where:
1) the apical lobe is proportionally shorter in L. exaratus
than in L. affinis; 2) it is also more tightly curled, nearly a
closed cylinder (vs. more open and expanded in L. affinis);
Male abdomen (Fig. 3B) narrow and long; somites 3–5
fused, sutures vaguely discernible; somite 6, median length
about 1.6 times that of telson. Distal half slightly broader
than proximal half, lateral margins slightly concave. Telson
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Fig. 4. Left G1. A–D, Leptodius exaratus (H. Milne Edwards, 1834): A, B, neotype, 23.7 × 16.1 mm (NHM 1881.10), Karachi, Pakistan;
C, 23.6 × 15.5 mm (ZRC 2012.0111), Qushm, Persian Gulf; D, 21.5 × 14.2 mm (MNHN-B6640), Nosy Be, Madagascar. E–I, Leptodius
affinis (De Haan, 1835): E, H, 25.9 mm × 16.6 mm, (NMST-Cr 6425), Shibasaki Hayama, Japan; F, 13.9 × 9.0 mm (ZRC 2012.0119), Con
Dau, Vietnam; G, 23.9 × 15.4 mm (ZRC 2012.0110), Tranquebar, S. E. India; I, 29.5 × 19.3 mm, (ZRC 2012.0117), Negros, Philippines.
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3) the angle formed between the apical lobe and the rest of
the G1 is larger (vs. less in L. affinis, giving it a more bent
appearance); and 4) the ventral lip of the apical lobe has fewer
outgrowths in L. exaratus than in L. affinis (for L. exaratus
see Forest & Guinot, 1961: 62, fig. 54; Serène, 1984: 180,
fig. 106; for L. affinis see Serène, 1962: 258, fig. 1; Yeh et
al., 2006: 73, fig. 2C, D).
1892; Buitendijk, 1960); China (Stimpson, 1907; Gordon,
1931; Shen, 1932, 1937; Dai et al., 1986; Dai & Yang, 1991);
Hong Kong (Stimpson, 1907); Philippines (Estampador, 1937,
1959; Buitendijk, 1960; Garth & Kim, 1983); Micronesia
(Miyake, 1939, 1940); Gilbert Is. (Holthuis, 1953); Thailand
(Buitendijk, 1960); Vietnam (Serène, 1962); Taiwan (Chang,
1963; Yeh et al., 2006).
The following discussion in this paper shows that these
records refer to L. affinis (De Haan, 1835), and that at least
some of the various synonyms in the literature are justified.
Concerning records of “L. exaratus” from the Hawaiian
islands in the central Pacific (viz. Stimpson, 1858; Rathbun,
1906; Edmondson, 1925, 1946, 1962; Titgen, 1987; Castro,
2011), it is clear after examining the published figures and
material collected from there that these are not L. exaratus or
L. affinis as defined at present, and belong to other species.
They will be treated and discussed elsewhere.
Central Pacific: Pacific (Dana, 1852); Samoa, Fiji, Caroline
Is. (Ortmann, 1893; Buitendijk, 1960); Nauru, Marshall Is.
(Balss, 1938); Tuamotu Archipelago (Holthuis, 1953); Palau
(Takeda, 1976); Marianas (Paulay et al., 2003).
South Pacific: New Caledonia (A. Milne-Edwards, 1873;
Takeda & Nunomura, 1976); French Polynesia (Boone,
1934).
DISCUSSION
Distribution. — Leptodius affinis (De Haan, 1835) is found
in the eastern Indian Ocean and in the western and central
Pacific Ocean, ranging from the eastern coast of India all the
way to the oceanic islands of French Polynesia; extending
northward to central Japan and southward to southwestern
and southeastern Australia. It has been recorded from the
following localities:
The taxonomy of one of the most common intertidal IndoWest Pacific xanthid crab species, Leptodius exaratus (H.
Milne Edwards, 1834) has generally been regarded as stable
and most recent syntheses do not question its identity (e.g.,
Serène, 1984). Following the general consensus, Ng et al.
(2008: 203) listed just four junior synonyms under L. exaratus:
Cancer inaequalis Olivier, 1791, Cancer inaequalis Audouin,
1826, Leptodius lividus Paul’son, 1875, and Xantho exaratus
var. typica Ortmann, 1893. However, Cancer inaequalis
Olivier, 1791, is now regarded as a species of Xanthodius
Stimpson, 1859, an American genus. This confusion probably
arose from material identified as Cancer inaequalis from the
Red Sea by Audouin (1826) but is today believed to be L.
exaratus instead (see Guinot & Cleva, 2009).
Eastern Indian Ocean: Mergui Archipelago (De Man, 1887a;
Alcock, 1898; Chopra & Das, 1937); Bay of Bengal (Alcock
& Anderson, 1894; Chopra & Das, 1937); Laccadive Sea
(Alcock & Anderson, 1894); Andamans (Alcock, 1898;
Sankarankutty, 1962); Gulf of Mannar (Gravely, 1927);
Myanmar (Alcock, 1898; Buitendijk, 1960); Penang (Alcock,
1898); Sri Lanka (Alcock, 1898; Laurie, 1906; Sankarankutty,
1966); western Thailand (Ng & Davie, 2002).
The problems associated with L. exaratus and allied taxa,
however, have been understated. Stimpson (1907) discussed
the taxonomy of Leptodius exaratus (as Chlorodius exaratus)
and commented that while he chose to recognise a number of
varieties, they also appeared to intergrade. He commented:
Western Pacific: Japan (De Haan, 1835; Krauss, 1843; Miers,
1879; Ortman, 1893; Stimpson, 1907; Balss, 1922; Yokoya,
1933; Sakai, 1934, 1936, 1939, 1965, 1976; Takeda & Miyake,
1976; Yamaguchi et al., 1976; Takeda, 1978; Yamaguchi &
Baba, 1993; Minemizu, 2000); Australia (Haswell, 1882;
Ortmann, 1893; Boone, 1934; Balss, 1935; McNeill, 1968;
Jones & Morgan, 1994; Davie, 2002, 2011; Poore, 2004);
Korea (Miers, 1879; Kim, 1970, 1973); Indonesia (De Man,
“… one would scarcely be prepared to find so much variety
in the character of the surface, the number and shape of the
lateral teeth, and the sculpture of the feet, as we see in the
Fig. 5. Results of morphometric analysis of Leptodius exaratus (H. Milne Edwards, 1834) (A–C: n= 28) and L. affinis (De Haan, 1835) (A,
B: n = 99; C: n = 64). Box plots depicting maxima, minima, median, and quartile values for: A, ratios of the sternite 4 width and combined
length of sternites 3 and 4; B, ratios of the frontal and carapace widths; C, ratios of the apical lobe length and total length of the G1.
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Stimpson (1907) compared his varieties to three other species
described by De Haan from Japan: Cancer (Xantho) affinis
(cf. De Haan, 1835: 48, pl. 13, fig. 6), Cancer (Xantho)
distinguendus (cf. De Haan, 1835: 48, pl. 13, fig. 7), and
Cancer (Xantho) lividus (cf. De Haan, 1835: 48, pl. 13, fig.
8). Cancer distinguendus is now in the genus Macromedaeus
Ward, 1942. Cancer (Xantho) lividus De Haan, 1835,
however, is a junior homonym of Cancer lividus Latreille,
in Milbert, 1812 (presently in the genus Juxtaxanthias Ward,
1942), although these species are clearly different taxa.
The name Cancer (Xantho) affinis De Haan, 1835, on the
other hand, seems to have been largely forgotten. Stimpson
(1907), however, did not make any firm decisions on the
validity of Cancer affinis nor of Cancer lividus. Ortmann
(1893) considered both Cancer (Xantho) affinis De Haan,
1835, and Cancer (Xantho) lividus De Haan, 1835, as junior
subjective synonyms of Xantho exaratus var. typica. This has
been followed by other authors, e.g., Buitendijk (1960), as
Xantho exaratus, and Barnard (1950), as Xantho (Leptodius)
hydrophilus. Likewise, Yamaguchi & Baba (1993: 446,
447, fig. 164A, B), who examined De Haan’s material in
Leiden and figured the types, regarded both names as junior
subjective synonyms of Leptodius exaratus, but without
comment. Interestingly, both of De Haan’s names have
generally been missed in most treatments of Japanese and
East Asian taxa (e.g., Sakai, 1965, 1976; Dai et al., 1986;
Dai & Yang, 1991; Ng et al. 2001, 2008).
present instance, these characters being in other genera and
species of the highest specific importance. The varieties
described below, however, are found to run into each other
in all the characters which at first sight strike the examiner
as specific, and several of them are often found living
together under circumstances which do not fail to impress
the collector with the idea that they are one and the same
species.” (Stimpson, 1907: 52)
He noted that Chlorodius sanguineus H. Milne Edwards,
1834, is probably not a good species and chose to only
recognise it as a variety of L. exaratus. In addition, he
recognised eight other varieties (here listed in the order
Stimpson named them): rugosus (Bonin Islands), pictus
(Simoda, Japan), latifrons (Loo Choo), typicus (Japan and
China), acutidens (Loo Choo), cupulifer (Bonin Islands), latus
(Hong Kong), and granulosus (Hong Kong). All these names
are nomenclaturally available as they were all named before
1960 (ICZN, 1999) and were accompanied by descriptions,
figures and/or comparisons. Bonin Islands are today known
as the Ogasawara Islands, while Loo Choo is the old name
for the Ryukyu Islands (particularly, Okinawa).
With regard to “Chlorodius exaratus var. typicus”, Castro
et al. (2004: 39) discussed the use of the epithet “typica”
for various species of Trapeziidae, either used as a form
or variety. It was a common practice in the late 19th and
early 20th centuries for taxonomists to name the typical or
nominate form or variety (nomenclaturally equivalent to the
modern subspecies) as “forma typica”. However, the Code
(1999) allows for all forms and varieties to be recognised as
available names equivalent to species if they were described
before 1960. The use of “var. typicus” for one of Stimpson’s
forms therefore makes this an available name. Ortmann
(1893) first used the name Xantho exaratus var. typica, so
this name has priority over Chlorodius exaratus var. typicus
Stimpson, 1907. Ortmann’s (1893) “var. typica” was based
on material he had from the West Pacific region, with only
one male specimen from the Red Sea; and he also listed De
Haan’s (1835) two taxa under his synonymy for the form.
Ortmann’s (1893) var. typica can therefore be referred to
either L. exaratus s. str. or L. affinis, depending on which
specimen is selected (see later).
Serène (1962) added to the confusion when he described a
new species from Vietnam, L. nigromaculatus. He noted that
his new species was different from typical L. exaratus from
the Indian Ocean, but that it was very close to the variety
“pictus” of Stimpson (1907) (Serène, 1962: 259). In fact,
he commented:
“Stimpson (1907) définit 9 variétés d’exaratus, réduisant
sanguineus au rang de variété d’exaratus; s’il laisse gracilis
comme espèce distincte, il écrit « qu‘elle n’est peut-être qu’une
variété lisse de exaratus ». Il pense toutefois que de futures
observations permettront sans doute « la reconnaissance de
caractères constants définissant des espèces vraies dans ce
qu’il ne considère encore que comme des variétés » Il est
possible que waialuanus Rathbun, 1906 soit la var. latus
Stimpson; australis Ward, 1939, la var. rugosus Stimpson;
leptodon Forest et Guinot, 1961, la var. acutidens ou la var.
cupulifer Stimpson, nigromaculatus nov. sp., la var. pictus
Stimpson.” (Serène, 1962: 260)
However, from his discussion, it is clear that he believed
that none of the new names proposed by Stimpson could
be used. He also did not discuss the validity of De Haan’s
two names noted above. He probably took the advice of
Barnard (1950) who remarked about L. exaratus (as Xantho
(Leptodius) hydrophilus):
“Stimpson (1907, Smiths. Misc. Coll., xlix, pp. 52 sqq., pl.
6, figs. 3, 4, 6–9) discusses this species and several varieties,
some of which have a “supplementary tooth” on the anterolateral margin, making 5 in all (as in quinquedentatus). At
that time the importance of male 1st gonopod as a specific
Fig. 6. Distribution map of material examined for this study. Legend:
triangles = Leptodius exaratus (black triangle, locality of neotype);
ovals = Leptodius affinis (type locality: Japan).
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Lee et al.: Taxonomy of Indo-West Pacific Leptodius exaratus
character was not realized, so that it is not only impossible
to recognize any of his varieties from the descriptions, but
it is also quite probable that several distinct species are
confused.” (Barnard, 1950: 225)
The remaining two varieties, var. pictus and var. typicus
cannot be effectively separated. Stimpson (1907; as well
as Ortmann, 1893) recognised a typical common form
in the Chinese and Japanese seas and other parts of the
Pacific Ocean, and this variety agrees very well with what
is recognised here as Leptodius affinis. As discussed above,
Serène (1962) had already commented that his species,
L. nigromaculatus, is very close to var. pictus, sharing a
smooth posterior carapace region, broad and non-projecting
triangular anterolateral teeth and the yellowish colour of the
carapace. As such, the oldest names available for this entire
group should be Cancer (Xantho) affinis De Haan, 1835,
and Cancer (Xantho) lividus De Haan, 1835. Both names are
regarded by the current Code as simultaneously published
as they appeared in the same publication, and seniority must
be chosen by First Reviser action. We hereby select Cancer
(Xantho) affinis De Haan, 1835, as having priority over
Cancer (Xantho) lividus De Haan, 1835, whenever the two
names are regarded as synonyms. This is mainly because
Cancer (Xantho) lividus De Haan, 1835, may be confused
with the unrelated Cancer lividus Latreille, in Milbert, 1812,
which is now in the genus Juxtaxanthias.
While this is true and Barnard’s observations are valid, the
fact remains that all of the new names proposed by Stimpson
(1907) are actually nomenclaturally available. And because of
their date, some may be senior synonyms of Leptodius species
names established since 1907, including L. nigromaculatus
Serène, 1962. The problem is compounded by the fact that
none of Stimpson’s specimens are extant (see Evans, 1967;
Deiss & Manning, 1981; Manning, 1993; Manning & Reed,
2006). The only solution is to examine a good series of
specimens in the area where Stimpson (1907) obtained his
specimens (East Asia), find topotypes when possible, match
them against his descriptions and figures, and designate the
appropriate neotypes to fix the identity of these names.
The present authors have gathered a large collection of
“Leptodius exaratus” from China, Japan, Korea, Taiwan, the
Philippines, Singapore and adjacent areas, and they believe
that all of Stimpson’s nominal varieties can be identified
from this collection. The “supplementary” anterolateral tooth
mentioned by Stimpson (1907) is, in fact, a reliable character
and not as variable as he had indicated. Two varieties have this
feature: var. rugosus and var. latus, both of which are allied to,
or synonymous with what is currently known as L. sanguineus
(H. Milne Edwards, 1834) and/or L. philippinensis Ward,
1941. Six varieties have four anterolateral teeth: var. pictus,
var. latifrons, var. typicus, var. acutidens, var. cupulifer, and
var. granulosus. On the basis of Stimpson’s (1907) description
and figure, var. granulosus, is likely to be synonymous
or very close to Macromedaeus distinguendus. Two taxa,
var. acutidens and var. cupulifer may be synonymous, and
because both have a strongly projecting front with acuminate
anterolateral teeth, they are unlikely to be L. exaratus or L.
affinis as presently defined. From Stimpson’s description
and figures, these two taxa are closely allied or synonymous
with L. davaoensis Ward, 1941, and/or L. leptodon Forest &
Guinot, 1961 (see Takeda, 1976). The two latter species were
regarded as synonyms by Takeda (1980), but comparisons by
Mendoza (2010) suggest they may be separate taxa. A third
Stimpson variety, latifrons, is probably also synonymous
with var. acutidens and var. cupulifer. According to his
description (no figure was provided), Stimpson’s (1907) var.
latifrons differs from var. typicus and var. pictus only in its
slightly wider front and angular carapace anterolateral teeth
and little else. His description and comments, however, are
far too brief to ascertain much. The present authors are of
the opinion that var. latifrons is unlikely to be var. typicus
nor var. pictus because the most common form present in
Okinawa (on the basis of the extensive material collected
and examined) is actually not L. affinis which has yet to be
collected from the island. In terms of the carapace and G1
morphology, these Okinawan specimens are much closer to
his var. acutidens/var. cupulifer, as well as L. davaoensis/L.
leptodon. All the other specimens examined share the same
carapace features as well as the same G1 structure and we
believe they are synonymous.
Four names: Xantho exaratus var. typica Ortmann, 1893,
Chlorodius exaratus var. typicus Stimpson, 1907, Chlorodius
exaratus var. pictus Stimpson, 1907, and Leptodius
nigromaculatus Serène, 1962, now become junior subjective
synonyms of Cancer (Xantho) affinis De Haan, 1835. To
stabilise the taxonomy of this species, it is necessary to select
a lectotype and neotypes for Ortmann’s (1893) and Stimpson’s
(1907) varieties, respectively. The lectotype of Cancer
(Xantho) affinis (a female, RMNH D 44644) is hereby selected
as also the simultaneous lectotype of Xantho exaratus var.
typica Ortmann, 1893, by reason of this De Haan name being
included in the synonymy of Ortmann’s “var. typica”, and in
accordance with Article 72.4 of the Code (1999) concerning
type series. Both names now become objective synonyms.
Ortmann (1893), in listing down various names in his
synonymy for Xantho exaratus var. typica (e.g., Chlorodius
exaratus H. Milne Edwards, Cancer (Xantho) affinis De Haan,
Cancer (Xantho) lividus De Haan, etc.), effectively used the
type material for these species as syntypes of his “var. typica”
by indication, therefore making them available for selection as
lectotypes by subsequent workers. The specimens examined
by Ortmann from East Asia (Ortmann, 1893: 446) are now
paralectotypes. The lectotype of C. (X.) affinis is also selected
as the simultaneous neotype of Chlorodius exaratus var.
typicus Stimpson, 1907, and Chlorodius exaratus var. pictus
Stimpson, 1907, in accordance with the Code (1999). While
Stimpson (1907) recognised several varieties, the present
authors find that these two varieties, at least, are conspecific
due to the lack of differences in the morphology (especially
that of the G1) of topotypic specimens examined in this study.
Since the type localities of Cancer (Xantho) affinis De Haan,
Chlorodius exaratus var. typicus Stimpson, and Chlorodius
exaratus var. pictus can be considered to be generally the
same (all within the vicinity of mainland Japan), they are
unlikely to be different species, hence one name-bearing
type is sufficient for the three names. Selecting separate
neotypes for Stimpson’s varieties does not serve the cause
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Fig. 7. Variation in live colouration of male Leptodius affinis (De Haan, 1835). A–G, Changi Beach Park, Singapore, (ZRC 2012.1232), H,
Espiritu Santo, Vanuatu (ZRC 2012.1233). A, 31.0 × 20.2 mm; B, 26.8 × 16.9; C, 24.0 × 15.1 mm; D, 12.5 × 7.8 mm; E, 12.0 × 7.6 mm;
F, 18.6 × 12.3 mm; G, 22 × 14.2 mm. H, 17.9 × 11.8 mm.
199
Lee et al.: Taxonomy of Indo-West Pacific Leptodius exaratus
Oldham, R.N., commanding. Series II, No. 14. Journal of the
Asiatic Society of Bengal, 63:141–185.
of nomenclatural stability for this species, especially since
the taxonomy has been rather confused. While the discovery
of cryptic species within this group in the future cannot be
discounted, the best available evidence does not suggest this.
As discussed, since Stimpson’s taxa are almost impossible
to ascertain with confidence, it is better to objectively fix
their identities through designation of appropriate neotypes
and move on with their taxonomy. As for Stimpson’s (1907)
remaining varieties, each of those have been matched by the
present authors with available topotypic material with the
help of his descriptions and illustrations, but these will be
discussed in a larger separate work dealing with a complete
revision of Leptodius.
Alcock, A.W., 1898. Materials for a carcinological fauna of India.
No. 3. The Brachyura Cyclometopa. Part 1. The Family
Xanthidae. Journal of the Asiatic Society of Bengal, 67(2/1):
67–233.
Audouin, V., 1826. Explication sommaire des planches de crustacés
de l’Egypte et de la Syrie, publiées par Jules-César Savigny,
membre de l’Institut; offrant un exposé de characteres naturels
des genres avec la distinction des especes. Description de
l’Egypte ou recueil des observations et des recherches qui ont
été faites en Egypte pendant l’expedition de l’armée francaise.
Histoire naturelle, 1(4): 77–98.
Balss, H., 1922. Östasiatische Decapoden. IV. Die Brachyrhynchen
(Cancridea). Archiv für Naturgeschichte, 88A(11): 94–166.
The colour of what has been called “Leptodius exaratus”
in East Asia is extremely variable and is not a reliable
distinguishing character (see Todd et al., 2009). The large
series of specimens examined for this study demonstrates
this (Fig.7).
Balss, H., 1924. Decapoden des Roten Meeres. III. Die Parthenopiden,
Cyclo- und Catometopen. In: Expedition S.M. Schiff “Pola” in
das Rote Meer, nördliche und südliche Hälfte 1895/96-1897/98.
Zoologische Ergebnisse. XXXIV. Denkschriften der kaiserlichen
Akademie der Wissenschaften zu Wien, Mathematischnaturwissenschaftliche Klasse, 99(6): 1–18.
Finally, it is important to state that the taxonomy of the genus
Leptodius is far from satisfactorily resolved (see Ng et al.,
2008). In addition to Stimpson’s varieties, several names, viz.
Chlorodius hombronii Lucas, in Lucas & Jacquinot, 1853,
Leptodius efferens Rathbun, 1906, Leptodius waialuanus
Rathbun, 1906 [the holotype of this species has been examined
by two of the authors (JCEM, PKLN), it actually belongs in
the genus Etisus H. Milne Edwards, 1834], Leptodius planus
Ward, 1934, Leptodius australis Ward, 1936, and the types
thereof (if extant) need to be re-examined in greater detail.
This will be part of a more substantial work, now in progress,
which will revise the entire genus.
Balss, H., 1935. Brachyura of the Hamburg Museum Expedition to
South-western Australia, 1905. Journal of the Royal Society of
Western Australia, 21: 113–151.
Balss, H., 1938. Die Dekapoda Brachyura von Dr. Sixten
Bocks Pazifik-Expedition, 1917–1918. Göteborgs Kungliga
Vetenskaps-och Vitterhets-Samhälles Handlingar, (B)5(7):
1–85, pls. 1, 2.
Barnard, K. H., 1950. Descriptive catalogue of South African
decapod Crustacea (crabs and shrimps). Annals of the South
African Museum, 38: 1–837.
Boone, L., 1934. Crustacea: Stomatopoda and Brachyura. Scientific
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Vanderbilt, Commanding. Bulletin of the Vanderbilt Marine
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ACKNOWLEDGEMENTS
Bouvier, E -L., 1915. Décapodes marcheurs (Reptantia) et
stomatopodes recueillis a l’île Maurice par M. Paul Carie.
Bulletin Scientifique de la France et de la Belgique, 48:178–
318.
The authors are most grateful to the following for their
help with specimen loans: Tomoyuki Komai (CBM), Laure
Corbari and Danièle Guinot (MNHN), Paul Clark (NHM),
Hironori Komatsu (NMST), and Tan Siong Kiat and Tan
Swee Hee (RMBR). The first author thanks the Raffles
Museum of Biodiversity Research and the Systematics &
Ecology Laboratory, National University of Singapore, for
allowing him to use their facilities during his visit for the
present study. This work was supported by a grant from the
Marine Biotechnology Programme funded by the Ministry of
Land, Transport Maritime Affairs of the Korean Government,
the Basic Science Research Program through the National
Research Foundation of Korea (NRF) funded by the Ministry
of Education, Science and Technology (2011-0009155),
and the Ministry of Land, Transport and Maritime Affairs
(PM56641).
Buitendijk, A. M., 1960. Biological results of the Snellius Expedition
XXI. Brachyura of the families Atelecyclidae and Xanthidae
(Part I). Temminckia, 5: 252–338.
Castro, P., P. K. L. Ng & S. T. Ahyong, 2004. Phylogeny and
systematics of the Trapeziidae Miers, 1886 (Crustacea:
Brachyura), with the description of a new subfamily. Zootaxa,
643: 1–70.
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(Crustacea: Decapoda: Anomura, Brachyura) of the Hawaiian
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of 19 new records. Tunghai Journal, 5: 95–118.
Chopra, B. N. & K. N. Das, 1937. Further notes on Crustacea
Decapoda in the Indian Museum. IX. On three collections of
crabs from Tavoy and Mergui Archipelago. Records of the
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