Mendoza et al.: Xanthidae from Christmas and Cocos Islands
RAFFLES BULLETIN OF ZOOLOGY Supplement No. 30: 274–300
Date of publication: 25 December 2014
http://zoobank.org/urn:lsid:zoobank.org:pub:7F45C9E6-39AC-4A14-9AE6-003E85DF1CD8
New rock crab records (Crustacea: Brachyura: Xanthidae) from
Christmas and Cocos (Keeling) Islands, Eastern Indian Ocean
Jose Christopher E. Mendoza1*, Robert M. Lasley Jr.1 & Peter K. L. Ng1,2
Abstract. The xanthid crab fauna of Christmas Island and the Cocos (Keeling) Islands, Australian territories in the
Indian Ocean, is documented. A total of 107 species of xanthid crabs are recorded, with 30 new species records from
either or both territories. Two potentially new species are included in this listing, and the taxonomy of Paractaea
rufopunctata plumosa Guinot, in Sakai, 1976, and Leptodius planus Ward, 1934, is discussed.
Key words. Christmas Island, Cocos (Keeling) Islands, Decapoda, Xanthidae, checklist, species distribution
INTRODUCTION
All material examined are deposited in the Zoological
Reference Collection (ZRC) of the Lee Kong Chian Natural
History Museum (formerly Rafles Museum of Biodiversity
Research), National University of Singapore. Measurements
are shown as maximum carapace width (CW) by median
carapace length (CL), in millimeters. The following
abbreviations are used: coll. – collected by; det. – determined
by; I./Is. – island/islands, respectively; juv. – juvenile;
ovig. – ovigerous; and stn – station. Terminology of the
carapace regions follows Serène (1984). Only new records
and those species requiring further comment are treated in
the systematic account. Synonymies are restricted to records
from Christmas & Cocos (Keeling) islands only, if available.
Where possible, colour photographs of whole animals are
provided if the live colouration is not known for that species.
Illustrations of the G1 are also provided for those species
where the G1 morphology is not, or poorly, known.
The brachyuran fauna of Christmas Island (CI) and the Cocos
(Keeling) Islands (CK) has been documented by previous
workers since the early 20th century (Calman, 1909; Wood
Jones, 1909; Balss, 1934; Ward, 1934; Tweedie, 1947, 1950;
George, 1978; Morgan, 2000; Davie, 2002; Orchard, 2012).
Collections from the recent expeditions to those islands by the
Rafles Museum of Biodiversity Research and the Queensland
Museum, with support from the Christmas Island National
Park and the relevant Australian authorities, have made it
possible to review this fauna. The present work focuses on
the xanthid crabs of these islands and provides an updated
checklist. The relevant literature containing species records
from Christmas Island and the Cocos (Keeling) Islands as
well as station records from the recent expeditions to those
islands in the years 2010–2012 are summarised in Table 1
for easy reference.
A total of 107 species comprise the combined xanthid crab
fauna (Xanthidae sensu Ng et al., 2008) of Christmas (83 spp.)
and the Cocos (Keeling) (59 spp.) islands, both Australian
territories in the eastern Indian Ocean (Table 1). Of these,
30 are newly recorded from either or both territories (CI =
23 spp.; CK = 11 spp.; CI+CK = 4 spp.); with at least two
species new to science, to be described elsewhere. All the
xanthid subfamilies except Antrocarcininae Ng & Chia,
1994, Garthiellinae Mendoza & Manuel-Santos, 2012,
Glyptoxanthinae Mendoza & Guinot, 2011, and Trichiinae
De Haan, 1841 (=Zalasiinae Serène, 1968) are represented.
A necessary note on the station names used in the material
examined listing in this paper: for simplicity and uniformity,
we have opted to use the alpha numeric codes beginning
with “CI1”, “CI2”, and “CI3” to denote Christmas Island
stations for the three consecutive trips in 2010, 2011, and
2012, respectively. Likewise, “CK1” and “CK2” are used
for Cocos (Keeling) stations for the years 2011 and 2012,
respectively. Field labels accompanying the actual specimens
may vary only in the year-related station preix, but not the
actual number (e.g., the ield label accompanying a specimen
from stn CI2-09, may have been written as “CI-09-2011”,
that is, station #09 in Christmas Island, collected in the
year 2011). A detailed account of the stations is provided
elsewhere in this special issue (see Tan et al., 2014).
1
Department of Biological Sciences, Faculty of Science, National University of
Singapore, 14 Science Drive 4, 117543 Singapore; Email: jcmendoza@nus.edu.sg
(*corresponding author JCEM)
2
Lee Kong Chian Natural History Museum, Faculty of Science National University
of Singapore, 6 Science Drive 2, 117546 Singapore
© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)
274
RAFFLES BULLETIN OF ZOOLOGY 2014
SYSTEMATIC ACCOUNT
Remarks. Guinot (1969) recognised and described seven
subgroups within Paractaea rufopunctata (H. Milne-Edwards,
1834), including the nominal subspecies, P. rufopunctata
rufopunctata (type locality: Mauritius), based on the setation
and sculpturing of the dorsal carapace regions, as well as
on their geographic distribution. The other six were named
as “formes”, viz. P. rufopunctata forme illusoria (Red Sea),
P. rufopunctata forme plumosa (Paciic), P. rufopunctata
forme primarathbunae (Hawaii), P. rufopunctata forme
tertiarathbunae (Hawaii), P. rufopunctata forme intermedia
(Bikini Atoll, Hawaii), P. rufopunctata forme africana
(eastern Atlantic) and P. rufopunctata forme nodosa (western
Atlantic), the last originally described by Stimpson (1860) as
Actaea nodosa. Serène (1984) also recognised and described
three additional “formes” from the western Indian Ocean,
namely: P. rufopunctata forme frontalis, P. rufopunctata
forme waltersi and P. rufopunctata forme sanctaeluciae. Two
of these “formes” were eventually recognised and formally
designated as subspecies – P. rufopunctata plumosa Guinot,
in Sakai, 1976, and P. rufopunctata africana Guinot, 1976.
Superfamily XANTHOIDEA MacLeay, 1838
Family XANTHIDAE MacLeay, 1838
Subfamily ActAeinAe Alcock, 1898
Actaeodes tomentosus (H. Milne-Edwards, 1834)
Actaea tomentosa, Calman, 1909: 705; Ward, 1934: 19; Tweedie,
1947: 27
Actaeodes tomentosus, Morgan, 2000: 121 (table); Davie, 2002: 513
Material examined. Cocos (Keeling) Is.: 1 ♂, 31.5 × 19.2
mm (ZRC), stn CK1-02.
Remarks. This species is widespread throughout the
Indo-West Paciic region (viz. Serène, 1984) and has been
reported from Christmas Island (Calman, 1909; Ward, 1934;
Morgan, 2000).
New CK record.
Ng et al. (2008: 195, 196, 207) considered most of the forms
of P. rufopunctata as unavailable names, citing Article 10.2
of the International Code for Zoological Nomenclature,
with the exception of P. nodosa (Stimpson, 1860) which
they raised to full species, and P. rufopunctata plumosa
and P. rufopunctata africana, which they considered valid
subspecies. Both subspecies are here recognised as full
species on account of the clear morphological distinctions
between them and the typical P. rufopunctata (viz. Guinot,
1969, 1976; Serène, 1984).
Gaillardiellus rueppelli (Krauss, 1843)
(Fig. 1A)
Material examined. Cocos (Keeling) Is.: 1 ♀, 20.1 × 14.6
mm (ZRC), stn CK2-09.
Remarks. This species, irst described from Natal, South
Africa (Krauss, 1843), is widespread throughout the IndoWest Paciic region (Guinot, 1976). A related species, G.
superciliaris, was recorded by Tweedie (1950) from the
Cocos (Keeling) Islands, but the present specimen can be
conidently excluded from that taxon. Serène’s (1984) key to
the genus separates it from species such as G. superciliaris
(Odhner, 1925) and G. alphonsi (Nobili, 1905) by the
presence of 4 (instead of 3) lobes on the anterolateral margin
of the carapace after the exorbital angle and the lack of
subdivision of the 3M region of the carapace. Furthermore,
the present specimen has relatively sparser and shorter setae
on the carapace and pereopods compared to G. superciliaris
(cf. Guinot, 1976: pl. 16 ig. 3). The present specimen can
also be distinguished from the similar G. orientalis (Odhner,
1925) by the morphological features of the carapace such as
the broad furrows separating the carapace regions, presence
of fewer and smaller granules, and absence of tufts of
plumose granules on any of the dorsal carapace regions
(versus narrower furrows, bigger and compacted granules,
and presence of tufts of plumose setae on 1M, 2M in G.
orientalis; viz. Serène, 1984).
Paractaea plumosa has been reported from Japan, Kiribati,
Tuvalu, Marutea, the Nicobars, and Madagascar (Guinot,
1969; Serène, 1984). The present specimens, all females,
match well with the description by Guinot (1969: 248, ig.
21) and Serène (1984: 124, pl. 26 ig. C), particularly in
the carapace having four anterolateral teeth apart from the
exorbital angle, a heart-shaped 5L areole, an undivided 1P,
and abundant tufts of long setae on the dorsal surface as
well as on the ambulatory legs.
New CI record.
Pseudoliomera lata (Borradaile, 1902)
(Fig. 1C)
Actaea lata, Balss, 1934: 226
Pseudoliomera lata, Morgan, 2000: 121 (table)
Material examined. Christmas I.: 1 ♀, 9.4 × 6.0 mm
(ZRC), stn CI3-17(085).
New CK record.
Remarks. This species was irst described from the Maldives
by Borradaile (1902), and has since been recorded also from
Japan (Serène, 1984). Pseudoliomera lata was listed by
Balss (1934) and Morgan (2000) as occurring in Christmas
Island, but was not illustrated or discussed. The present
specimen agrees well with the key to the genus by Serène
(1984), particularly in the areolation of the carapace and the
Paractaea plumosa Guinot, in Sakai, 1976
(Fig. 1B)
Material examined. Christmas I.: 1 ♀, 22.9 × 15.2 mm
(ZRC), stn CI1-D02; 1 ♀, 22.1 × 15.0 mm (ZRC), stn
CI2-D03(091); 1 ♀, 13.7 × 9.5 mm (ZRC), stn CI2-D15(171).
275
Mendoza et al.: Xanthidae from Christmas and Cocos Islands
Fig. 1. Live colouration. A, Gaillardiellus rueppellii (Krauss, 1843), 1 ♀, 20.1 × 14.6 mm (ZRC), stn CK2-09; B, Paractaea plumosa
Guinot, in Sakai, 1976, 1 ♀, 22.1 × 15.0 mm (ZRC), stn CI2-D03(091); C, Pseudoliomera lata (Borradaile, 1902), 1 ♀, 9.4 × 6.0 mm
(ZRC), stn CI3-17(085); D, Pseudoliomera violacea (A. Milne-Edwards, 1873), 1 ovig. ♀, 18.5 × 12.5 mm (ZRC), stn CI3-D05; E,
Chlorodiella cytherea (Dana, 1852), 1 ♂, 12.7 × 8.0 mm (ZRC, RL 17), stn CI3-17(087); F, Cyclodius drachi (Guinot, 1964), 1 ♂, 9.8
× 6.6 mm (ZRC, RL 25), stn CI2-D03.
276
RAFFLES BULLETIN OF ZOOLOGY 2014
presence of a comb-like row of stiff setae on the dactylus
of the irst ambulatory leg.
Islands, and has subsequently been reported from the
Hawaiian Islands to Madagascar. In external morphology,
C. cytherea is almost indistinguishable from C. davaoensis
Ward, 1941 and C. crispipleopa Dai, Yang, Song &
Chen, 1986. Dai et al. (1986: 342) provided a table of the
distinguishing features of C. crispileopa and C. cytherea: i.e.,
carapace, front, second anterolateral tooth, and G1. However,
examination of the extensive collection of this species from
Christmas Island indicates that these features are variable.
Even the G1, which is useful for identiication of other
chlorodielline species, varies from the distally “curled” C.
crispipleopa form (e.g. RL 1) (Dai & Yang, 1991: 341, ig.
169(1)) to the distally “hook-shaped” C. cytherea form (e.g.,
RL 6, RL 17) (Dai & Yang, 1991: 341, ig. 169(4)). A G1
igure of C. davaoensis was not provided by Ward (1941) or
since the original description. It is unclear whether these three
species are distinct. Examination of the type specimens and
material from the type localities is necessary to resolve this
problem, and a systematic revision of this species complex
and the Chlorodiellinae is being undertaken (RML & PKLN,
in prep.). For the moment, the earliest name, C. cytherea,
is attributed to the specimens reported here as they it the
original description of Dana (1852).
Pseudoliomera violacea (A. Milne-Edwards, 1873)
(Fig. 1D)
Material examined. Christmas I.: 1 ♂, crushed carapace,
not measured (ZRC), stn CI2-D17; 1 ovig. ♀, 18.5 × 12.5
mm (ZRC), stn CI3-D05.
Remarks. This rarely encountered species was irst described
as Lophactaea violacea by A. Milne-Edwards (1873), from
specimens from New Caledonia. It has since been recorded
by Buitendijk (1941) from Ambon, Indonesia. There have
been no subsequent records of this species elsewhere.
Pseudoliomera violacea is separated from other congeners
by its purplish colouration, the cristate anterolateral margins
of the carapace, and the wide, setae-illed, furrows between
the dorsal carapace regions (cf. Serène, 1984).
New CI record.
Subfamily chlorodiellinAe Ng & Holthuis, 2007
Chlorodiella cytherea Dana, 1852
(Fig. 1E)
New CI, CK record.
Cyclodius drachi (Guinot, 1964)
(Fig. 1F)
Material examined. Christmas I.: 1 ♂, 8.8 × 5.7 mm
(ZRC, RL 2) stn CI2-17; 2 ♀, 10.5 × 6.3 mm – 8.4 × 5.3
mm (ZRC, RL 4), stn CI1-31; 4 ♂, 11.1 × 6.9 mm – 10.3
× 6.3 mm, 1 ♀, 10.5 × 6.2 mm (ZRC, RL 5), stn CI2-22; 2
♂, 8.2 × 5.2 mm – 6.8 × 4.3 mm, 2 ♀, 6.7 × 4.2 mm – 5.2
× 3.4 mm (ZRC, RL 6), stn CI2-17; 1 ♂, 4.8 × 3.1 (ZRC,
RL 7), stn CI1-31; 1 ♂, 7.0 × 4.8 mm (ZRC, RL 8), stn
CI2-09; 1 ovig. ♀, 8.3 × 5.0 mm (ZRC, RL 9), stn CI1-03;
1 ♂, 8.3 × 5.1 mm, 1 ♀, 8.1 × 4.9 mm (ZRC, RL 10), stn
CI3-25; 1 ♀, 7.7 × 4.8 mm (ZRC, RL 12), stn CI3-17; 2 ♂,
9.1 × 5.8 mm – 9.5 × 6.3 mm, 1 ♀, 6.0 × 3.8 mm (ZRC,
RL 13), stn CI3-14; 1 ♂, 8.2 × 5.1 mm (ZRC, RL 14), stn
CI3-15; 1 ♀, 8.1 × 4.9 mm (ZRC, RL 15), stn CI3-14; 1 ♂,
9.3 × 6.2 mm (ZRC, RL 16), stn CI3-14; 1 ♂, 12.7 × 8.0
mm (ZRC, RL 17), stn CI3-17; 1 ♀, 4.1 × 2.7 mm (ZRC,
RL 18), stn CI3-13; 2 ♂, 7.2 × 4.5 mm – 7.2 × 4.4 mm,
1 ♀, 9.1 × 5.9 mm (ZRC, RL 19), stn CI3-16; 1 ♂, 5.8 ×
3.7 mm, 1 ♀, 8.1 × 5.2 mm (ZRC, RL 21), stn CI3-17; 3
♂, 4.3 × 2.7 – 10.2 × 6.2 mm, 2 ♀, 6.8 × 4.3 – 7.6 × 5.0
(ZRC, RL 23), stn CI3-15; 5 ♂, 5.7 × 3.6 mm – 10.0 × 6.0
mm, 5 ♀, 6.3 × 3.7 mm – 7.9 × 4.9 mm (ZRC), stn CI3-23.
Cocos (Keeling) Is.: 1 ♂, 12.0 × 7.4 mm (ZRC, RL 1),
stn CK1-08; 1 ♂, 6.9 × 4.4 mm, 1 ♀, 5.5 × 3.4 mm (ZRC,
RL 3) stn CK1-08; 1 ♂, 11.5 × 7.1 mm (ZRC, RL 11), stn
CK2-21; 1 ♂, 5.0 × 3.1 mm, 1 ♀, 8.8 × 5.6 (ZRC, RL 20),
stn CK2-21; 5 ♂, 9.0 × 5.6 mm – 11.3 × 7.0 mm, 2 ♀,
8.0 × 5.1 mm – 9.8 × 6.0 mm (ZRC, RL 22), stn CK2-21.
Material examined. Christmas I.: 1 ♂, 9.8 × 6.6 mm (ZRC
2013.1630), stn CI2-D03.
Remarks. This relatively rare chlorodielline species was
described from Madagascar and recorded from the Red Sea
by Guinot (1964). It has subsequently been reported from
Kenya, Mayotte and the Paracel (Xisha) Islands (Serène,
1984; Dai et al., 1986). Cyclodius drachi is most similar to
C. nitidus, especially with respect to carapace morphology.
Both are relatively wide and smooth, and have regions
defined by deep areolas, making identification difficult.
Beyond these features, several authors have differentiated
between them based on carapace granulation, division of 2M,
granulation of the chelipeds (especially in females), and the
shape of the male abdomen (Guinot, 1964; Serène, 1984;
Dai et al., 1986). However, these characters are variable
in C. drachi or dificult to discern, making examination
of the male gonopod necessary for accurate identiication.
The G1 of C. drachi is adorned distally with long, plumose
setae and has a spatulate tip (versus long, simple setae and
a tubular tip in C. nitidus) (Serène, 1984: igs. 47, 50). The
individual collected from Christmas Island is small and
has an underdeveloped G1. Its identity was veriied via a
comparison of the barcoding region of the mitochondrial
gene, COI, between our specimen and several large adults
from various localities (RML, unpublished data).
Remarks. Chlorodiella cytherea was described by Dana
(1852) from Tuamotu Archipelago, Tahiti and the Hawaiian
New CI record.
277
Mendoza et al.: Xanthidae from Christmas and Cocos Islands
Pilodius lavus Rathbun, 1894
Cymo cerasma Morgan, 1990
(Figs. 2A, 6C, D)
Material examined. Christmas I.: 1 ♂, 9.4 × 6.4 mm (ZRC,
RL 24), stn CI1-D17.
Material examined. Christmas I.: 1 ♂, 10.6 × 9.6 mm
(ZRC), stn CI1-D04 (w/ colour photo).
Cocos (Keeling) Is.: 1 ♂, 9.6 × 8.6 mm, 2 ♀, 7.5 × 7.0
mm, 11.8 × 10.6 mm (ZRC), CK1-08 (♂ w/ colour photo).
Remarks. As presently defined, Pilodius flavus (type
locality: Hawaiian Islands) has a widespread distribution
from the Hawaiian Islands to Madagascar, although it has
not been reported from the Red Sea. Clark & Galil (1993)
synonymised Chlorodopsis hawaiiensis Edmondson, 1962
and C. melanospinis Rathbun, 1911, with P. lavus, after
examining a large series of specimens including types of
P. lavus and C. melanospinis, and material identiied by
Edmondson (1962) as C. hawaiiensis. They concluded that
the external morphological differences between P. lavus
and C. melanospinis “appear to be just variation” and stated
that the G1s are identical (Clark & Galil, 1993: 1132). The
authors did not elaborate on their decision to synonymise
C. melanospinis under P. lavus beyond stating that material
identiied as C. hawaiiensis by Edmondson (1962) “proved
to be P. lavus.” However, Edmondson’s (1962: 225–226
igs. 21e, 22b) illustrations of the G1s of P. lavus and C.
hawaiiensis, although simplistic, differ from the G1 illustrated
by Clark & Galil (1993: 1164, ig. 4D–G). Close scrutiny of
numerous specimens indicates there are indeed different G1
morphologies (RML, pers. obs.). Furthermore, COI sequence
data from a widespread sample of specimens indicates two
divergent “P. lavus” clades (RML, unpublished data). There
are likely two, or possibly three, species lumped in P. lavus.
Re-examination of the types and other material examined
by these authors is necessary to stabilise the taxonomy of
the group and re-determine distributions. The one specimen
recorded here from Christmas Island has a G1 like that
illustrated by Edmondson (1962: 226, ig. 22b) as P. lavus.
Remarks. This species was described by Morgan (1990:
43, fig. 5) from northwestern Australia, and was also
mentioned by the same author to be present in Madang,
Papua New Guinea although no specimens from there was
expressly included among the material examined. The present
specimens have the G1 morphology (Fig. 6C, D) typical of
the species (cf. Morgan, 1990: ig. 5i, j). Some variation,
however, was noted between the CI and CK forms. While the
CI specimen agrees well with the illustration and description
by Morgan (1990), the CK specimens appear to be more
spinose on the carapace, chelipeds and ambulatory legs.
Furthermore, the CK specimens are similar to C. andreossyi
except for the G1 morphology and that the ingers of chelae
are pigmented, not white (but pigment does not extend into
palm in the male, as in C. melanodactylus).
New CI, CK record.
Cymo quadrilobatus Miers, 1884
(Fig. 2B)
Cymo quadrilobatus, Tweedie, 1950: 123; Davie, 2002: 528
Material examined. Christmas I.: 1 ♂, 12.2 × 11.4 mm
(ZRC) stn CI1-D14(126); 1 ♂, 16.7 × 15.5 mm (ZRC), stn
CI1-D17(165); 1 ♀, 17.8 × 16.5 mm (ZRC) stn CI1-D17
(164); 4 ovig. ♀, 12.7 × 11.5 mm – 17.5 × 16.0 mm (ZRC),
stn CI1-D18/19.
Cocos (Keeling) Is.: 2 ♀, 8.8 × 8.6 mm, 11.7 × 11.3 mm
(ZRC), stn CK2-12.
New CI record.
Subfamily cymoinAe Alcock, 1898
Remarks. This species was originally described from the
Red Sea (Miers, 1884), and has been reported from several
localities in the Indo-West Pacific region (viz. Serène,
1984). It was previously reported by Tweedie (1950) from
the Cocos (Keeling) Is.
Cymo andreossyi (Audouin, 1826)
Cymo andreossyi, Tweedie, 1950: 123
Material examined. Christmas I.: 1 ♂, 6.3 × 5.4 mm,
detached carapace (ZRC), stn CI3-16.
New CI record.
Remarks. Cymo andreossyi (type locality: Red Sea) has a
wide distribution and has been recorded in the South Africa,
Japan, Australia, and the Paciic and Western Indian oceans
(Serène, 1984). The present specimen, although partially
damaged, agrees well with the key and illustrations in Serène
(1984: 32, ig. 7, pl. 2 ig. C). It can be distinguished from
the similar C. melanodactylus De Haan, 1833, by the entirely
white ingers on both chelae and by the form of the G1 (cf.
Serène, 1984: ig. 8, pl.2 ig. B). This species was previously
recorded by Tweedie (1950) from the Cococs (Keeling) Is.
Subfamily etisinAe Ortmann, 1893
Etisus demani Odhner, 1925
(Figs. 2C, 6E)
Etisus demani, Tweedie, 1947: 31; Morgan, 2000: 122 (table);
Davie, 2002: 530
Material examined. Christmas I.: 1 ♂, 8.5 × 6.1 mm (ZRC),
stn CI2-22; 1 ♂, 22.0 × 14.6 mm (ZRC), 1 ♂, 28.5 × 18.9,
2 ♀, 11.8 × 8.2 mm, 17.5 × 12.0 mm (ZRC) stn CI3-16.
Cocos (Keeling) Is.: 1 ♂, 15.7 × 10.6 mm (ZRC), stn CK1-18.
New CI record.
278
RAFFLES BULLETIN OF ZOOLOGY 2014
Fig. 2. Live colouration. A, Cymo cerasma Morgan, 1990, 1 ♂, 10.6 × 9.6 mm (ZRC), stn CI1-D04; B, Cymo quadrilobatus Miers, 1884,
1 ♂, 16.7 × 15.5 mm (ZRC), stn CI1-D17(165); C, Etisus demani Odhner, 1925, 1 ♂, 28.5 × 18.9 (ZRC) stn CI3-16(071); D, Etisus
dentatus (Herbst, 1785), 1 juv. ♂, 19.4 × 14.4 mm (ZRC), stn CI3-16(072); E, Etisus odhneri Takeda, 1971, 1 ♂, 11.2 × 8.0 mm (ZRC),
stn CI1-D17(158); F, Etisus splendidus Rathbun, 1906, 1 ♀, 45.8 × 32.7 mm (ZRC), stn CI1-D17(169).
279
Mendoza et al.: Xanthidae from Christmas and Cocos Islands
Remarks. This species, originally described from Samoa
(see Odhner, 1925), has been previously reported from
Christmas Island (Tweedie, 1947; Morgan, 2000). It is
a wide-ranging species, having been recorded from the
Red Sea and the western Indian Ocean to the Hawaiian
Islands and the Tuamotus. The present specimens are from
Christmas and the Cocos (Keeling) islands. The G1 of the
CI and CK specimens (Fig. 6E) is similar in morphology to
that illustrated for E. demani by Takeda (1971: ig. 3C, D),
based on a specimen of unspeciied provenance, originally
deposited at the Zoological Laboratory, Kyushu University
(now transferred to the Kitakyushu Museum of Natural
History & Human History). The very slender tip of the G1
has a long apical lobe, where the free border is lined with
small, T-shaped evaginations. In contrast, the G1 illustrated
by Serène (1984: fig. 140), based on a specimen from
Madagascar, has a much shorter apical lobe, without the
T-shaped evaginations on the border (which is entire instead).
advanced than lateral in E. electra). The present specimen
agrees well with illustrations by Guinot (1964: ig. 23, pl. 5
ig. 2) and Serène (1984: pl. 31 ig. E). Also, Guinot (1964)
provided additional characters in the carapace supraorbital
margin, chelipeds, and G1 that can be used to differentiate
the two species.
New CK record.
Etisus odhneri Takeda, 1971
(Figs. 2E, 6F)
Material examined. Christmas Is: 1 ♂, 11.2 × 8.0 mm
(ZRC), stn CI1-D17(158); 1 ♂, 10.4 × 7.4 mm (ZRC), stn
CI1-D17(160); 1 ♀, 7.0 × 5.1 mm (ZRC), stn CI2-D12.
Cocos (Keeling) Is.: 1 ♂, 12.5 × 8.8 mm (ZRC), stn CK2-03.
Remarks. This species was described from Palau by Takeda
(1971), and has also been reported from coast of Kenya
(Serène, 1984). It can be distinguished from the similar E.
demani by the less prominent dorsal carapace areolae, the
absence of prominent intercalating denticles between the
teeth of the anterolateral margins of the carapace, and by
the G1 morphology (see Fig. 6F).
New CK record.
Etisus dentatus (Herbst, 1785)
(Fig. 2D)
Etisus dentatus, Tweedie, 1950: 120; Davie, 2002: 530
New CI, CK record.
Material examined. Christmas I.: 1 ♂, 72.0 × 49.6 mm
(ZRC), stn CI3-14(028); 1 ♂, 57.3 × 40.4 mm (ZRC), stn
CI3-14; 1 juv. ♂, 19.4 × 14.4 mm (ZRC), stn CI3-16(072).
Cocos (Keeling) Is.: 1 juv. ♂, 26.3 × 18.9 mm (ZRC), stn
CK2-21.
Etisus splendidus Rathbun, 1906
(Fig. 2F)
Material examined. Christmas I.: 1 ♀, 45.8 × 32.7 mm
(ZRC), stn CI1-D17(169).
Remarks. Etisus dentatus is a widespread Indo-Pacific
species, occurring from the Red Sea and western Indian
Ocean all the way to the Hawaiian Islands (Serène, 1984).
It belongs to that subgroup of Etisus spp. that are large
with a smooth, reddish carapace and distinct, curved teeth,
interspersed with smaller teeth, on the carapace anterolateral
margin, and with spinous ambulatory propodi and dactyli.
This species was previously recorded by Tweedie (1950)
from the Cocos (Keeling) Islands. The juvenile form is
igured here (Fig. 2D).
Remarks. This large species was irst described from the
Hawaiian Islands by Rathbun (1906) and has been recorded
throughout the Indo-West Paciic region from the Red Sea to
the central Paciic Islands (Serène, 1984). It can readily be
distinguished from the similar E. dentatus by the presence
of a low double-crest, bearing low, blunt spines, on the
superior margin of the cheliped palm, and the presence of
two (not one), subequal spines on the inner angle of the
cheliped carpus.
New CI record.
New CI record.
Etisus frontalis Dana, 1852
(Fig. 6A)
Subfamily euxAnthinAe Alcock, 1898
Euxanthus aff. exsculptus
Material examined. Cocos (Keeling) Is.: 1 ♀, 8.2 × 5.8
mm (ZRC), stn CK1-18.
Euxanthus exsculptus, Tweedie, 1950: 115, ig. 2a; Davie, 2002:
543 (in part). Not Cancer exsculptus Herbst, 1790
Remarks. This species was originally described from the
Sulu Archipelago in the Philippines (Dana, 1852) and has
also been recorded with certainty from Aldabra and Hikueru
(Tuamotu Archipelago) (Guinot, 1964; Serène, 1984). The
sole specimen reported here (Fig. 6A), a female, can be
distinguished from the similar (and co-occurring) E. electra
mainly by the less distinctly quadridentate front, and also
by having the mesial and lateral teeth of the frontal lobes
more level (versus strongly quadridentate, mesial teeth more
Material examined. Cocos (Keeling) Is.: 5 ♂, 35.5 × 23.6
mm – 52.2 × 34.0 mm, 3 ♀, 31.9 × 21.1 – 51.6 × 34.2 mm
(ZRC 1965.11.9.34-41), Pulo Cheplok, coll. C. Gibson-Hill,
1941, Balss det. as E. exsculptus.
Remarks. This species was not collected during the three
recent expeditions (2010–2012) but was recorded from
Cocos (Keeling) Is. by Tweedie (1950) as Euxanthus
280
RAFFLES BULLETIN OF ZOOLOGY 2014
Fig. 3. Live colouration. A, Medaeus elegans A. Milne-Edwards, 1867, 1 ovig. ♀, 10.6 × 7.4 mm (ZRC), stn CI3-D01; B, Paramedaeus
sp., 1 ♂, 21.7 × 13.1 mm (ZRC), stn CI3-D08(154); C, Liomera stimpsonii (A. Milne-Edwards, 1865), 1 ♂, 15.5 × 8.9 mm (ZRC), stn
CI2-D03(094); D, Liomera virgata (Rathbun, 1906), 1 ♂, 14.6 × 8.2 mm (ZRC), stn CI3-D02; E, Lybia leptochelis (Zehntner, 1894), 1
♂, 5.1 × 3.6 mm (ZRC), stn CI2-D08(132); F, Lachnopodus ponapensis (Rathbun, 1907), 1 ♂, 18.0 × 12.1 mm (ZRC) stn CI3-23.
281
Mendoza et al.: Xanthidae from Christmas and Cocos Islands
Subfamily liomerinAe Sakai, 1976
exsculptus (Herbst, 1790), based on material from the
extensive collections of C.A. Gibson-Hill. Examination of
these same specimens by one of the present authors (JCEM),
however, has revealed some variation in the carapace and
G1 morphology from that described for E. exsculptus sensu
stricto (lectotype designated by K. Sakai, 1999). The Cocos
(Keeling) Is. material, along with specimens from other
localities in the Indian Ocean will be described as a distinct
species in a larger work focusing on the revision of the
subfamily Euxanthinae (JCEM & PKLN, in prep.).
Liomera stimpsonii (A. Milne-Edwards, 1865)
(Fig. 3C)
Carpilodes stimpsonii, Tweedie, 1950: 113
Liomera stimpsoni, Davie, 2002: 544
Material examined. Christmas I.: 1 ♂, 15.5 × 8.9 mm
(ZRC), stn CI2-D03(094); 1 ♀, 7.9 × 4.6 mm, 2 juv., 4.7
× 3.1 mm, 4.8 × 3.2 mm (ZRC), stn CI2-D15; 1♂, 12.0 ×
7.0 mm, 1 ♀, 17.4 × 10.3 mm (ZRC), stn CI2-D18(196);
2♂, 11.9 × 7.0 mm, 14.2 × 8.1 mm, 1 ♀, 12.1 × 7.2 mm,
1 ovig. ♀, 13.2 × 7.7 mm (ZRC), stn CI3-D05.
Medaeus elegans A. Milne-Edwards, 1867
(Figs. 3A, 7E)
Material examined. Christmas I.: 1 ovig. ♀, 10.6 × 7.4
mm (ZRC), stn CI3-D01.
Cocos (Keeling) Is.: 1 ♂, 13.0 × 8.9 mm (ZRC), stn CK212; 1 ♂, 12.3 × 8.2 mm (ZRC), stn CK2-13
Remarks. This species was previously recorded from the
Cocos (Keeling) Islands by Tweedie (1950).
New CI record.
Remarks. This species was irst described by A. MilneEdwards (1867) from New Caledonia. It has since been
reported from the Hawaiian Islands, Gilbert Islands, Marshall
Islands, and the Philippines (Edmondson, 1962; Guinot, 1967;
Garth et al., 1987; Mendoza & Ng, 2010). Mendoza & Ng
(2010) commented that this taxon needs to be revised as it
is likely to represent a species complex. With more material
available from other localities throughout the Indo-West
Paciic region, it is now possible to elucidate the taxonomy
of Medaeus. The revision of this genus is part of a larger
work focusing on the revision of the subfamily Euxanthinae
(JCEM & PKLN, in prep.). The present specimens constitute
the irst record of this species from the Indian Ocean.
Liomera virgata (Rathbun, 1906)
(Figs. 3D, 6G–I)
Material examined. Christmas I.: 1 ovig. ♀, 10.8 × 6.2
mm (ZRC), stn CI2-D17; 1 ♂, 14.6 × 8.2 mm (ZRC), stn
CI3-D02.
Remarks. The present specimens agree well with the
description and igure of the type, collected from Au’au
Channel, Hawaiian Islands (cf. Rathbun, 1906: 843, pl. 8 ig.
3). This species has also been reported from the Amirante
Islands and Holothuria Bank, South China Sea (Serène,
1984). The live colouration (Fig. 3D) and the G1 morphology
(Fig. 6G–I) for this species are reported for the irst time.
New CI, CK record.
New CI Record.
Paramedaeus sp.
(Fig. 3B)
Subfamily PolydectinAe Dana, 1851
Material examined. Christmas I.: 1 ♂, 21.7 × 13.1 mm
(ZRC), stn CI3-D08(154).
Lybia leptochelis (Zehntner, 1894)
(Fig. 3E)
Remarks. A single male specimen was collected from
Christmas Island. The carapace morphology is starkly
different from other Paramedaeus species reported from
the Indian Ocean (e.g., P. octogesimus Ng & Clark, 2002,
P. simplex (A. Milne-Edwards, 1873)), particularly in the
dentition of the anterolateral margin of the carapace. While in
those species the four anterolateral teeth are acutely triangular
and well separated from each other by V-shaped troughs,
the CI specimen has more obtuse teeth barely separated by
narrow clefts. The specimen was collected from beneath a
limestone rock about 30 m from the entrance to Thundercliff
Cave. As the cavern bent almost 90 degrees just after this
entrance, the habitat was quite dark even at daytime. This
specimen has been found to be conspeciic with another
male from Guam I., and both will be described as new in a
forthcoming revision of the subfamily Euxanthinae (JCEM
& PKLN, in prep.).
Material examined. Christmas I.: 1 ♂, 5.1 × 3.6 mm
(ZRC), stn CI2-D08(132).
Remarks. The present specimen agrees well with the
descriptions and igures in Guinot (1976: 71, igs. 16C, 21E,
F, 22C, pl. 2 ig. 2) and Mendoza & Ng (2011: 52, ig. 1A).
There are nine teeth on each of the ingers of the chelae,
although the most distal tooth is much smaller than the rest.
The second and third anterolateral teeth are pronounced and
distinctly angular, and there is a tiny, vestigial tooth behind
the third anterolateral tooth. This species has been reported
from Ambon, Indonesia (type locality), Mozambique (Serène,
1984) and the Philippines (Mendoza & Ng, 2011).
New CI record.
New CI record.
282
RAFFLES BULLETIN OF ZOOLOGY 2014
Fig. 4. Live colouration. A, Leptodius planus Ward, 1934, 1 ♀, 10.2 × 7.0 mm (ZRC), stn CK2-21; B, Macromedaeus crassimanus (A.
Milne-Edwards, 1867), 1 ♂, 32.8 × 21.3 mm (ZRC), stn CI3-15; C, Nanocassiope alcocki (Rathbun, 1902), 1 ♀, 9.8 × 6.5 mm (ZRC),
stn CI3-D08(157); D, Paraxanthias aff. elegans, 1 ♂, 7.5 × 5.1 mm (ZRC), stn CI3-D02; E, Xanthias cherbonnieri Guinot, 1964, 1 ♂,
4.5 × 3.0 mm (ZRC), stn CI2-D18; F, Platypodia anaglypta (Heller, 1861), 1 ♂, 13.6 × 9.0 mm (ZRC), stn CI3-15(050).
283
Mendoza et al.: Xanthidae from Christmas and Cocos Islands
Subfamily xAnthinAe MacLeay, 1838
with the immediately preceding portion of the G1 (cf. Forest
& Guinot, 1961: ig. 58; Serène, 1984: ig. 107), whereas in
L. planus, the apical lobe forms a small curve distinct from
the curvature of the immediately preceding portion of the
G1 (Fig. 7A). Furthermore, the teeth on the anterolateral
margin of the carapace are broader and more uniform in
L. planus. The records of “L. gracilis” by Tweedie (1950)
and Morgan (2000) from Cocos (Keeling) and Christmas
islands, respectively, are here considered to be L. planus.
Lachnopodus ponapensis (Rathbun, 1907)
(Figs. 3F, 7B–D)
Paraxanthias haematostictus Ward, 1934: 20, pl. 2 ig. 3, 3a
Paraxanthias ponapensis, Tweedie, 1947: 29
Lachnopodus ponapensis, Morgan, 2000: 121 (table); Davie,
2002: 550
Material examined. Christmas I.: 1 ♂, 8.3 × 5.7 mm (ZRC),
stn CI1-13; 1 ♂, 15.4 × 10.4 mm (ZRC), stn CI2-09 [024];
1 ♂, 8.7 × 6.1 mm (ZRC), stn CI2-17 [075]; 3 ♂, 11.0 ×
7.7 mm – 29.0 × 12.7 mm, 2 ♀, 11.4 × 8.0 mm, 16.0 ×
10.9 mm (ZRC), stn CI2-22; 1 ♂, 18.0 × 12.1 mm (ZRC)
stn CI3-23(095), 1 ♂, 18.6 × 13.0 mm (ZRC), stn CI3-23;
1 ♂, 7.5 × 5.3 mm (ZRC), stn CI3-25.
Macromedaeus crassimanus (A. Milne-Edwards, 1867)
(Fig. 4B)
Material examined. Christmas I.: 1 ♂, 32.8 × 21.3 mm
(ZRC), stn CI3-15; 1 ♀, 26.8 × 17.8 mm (ZRC), stn CI3-16.
Remarks. This species was originally described from New
Caledonia (A. Milne-Edwards, 1867), and has a widespread
distribution, from Hawaii, Samoa and Tahiti all the way to
the Red Sea (Dai & Yang, 1991). The G1 of the present
specimen differs from that illustrated for the smaller male
syntype (MNHN; 28 × 18 mm) (cf. Serène, 1984: ig. 103)
in having a more extended and tapered distal tip (versus
bluntly rounded). It is more similar to the G1 illustrated
by Dai & Yang (1991: ig. 151(2)) from a male from the
Paracel (Xisha) Is., South China Sea. In all other respects,
the present specimens agree well with the description and
illustrations of M. crassimanus.
Remarks. This species was irst recorded from Christmas
Island as Paraxanthias haematostictus, a new species
described by Ward (1934). It was later synonymised under
Paraxanthias ponapensis (Rathbun, 1907) (type locality:
Ponape, Caroline Islands) by Tweedie (1947) after careful
comparison of the types of both species. Morgan (2000)
relected the latest accepted assignment of this species to the
genus Lachnopodus Stimpson, 1858. This species is most
readily distinguished from other species of Lachnopodus by
its distinctly spotted carapace (Fig. 3F). The G1 is illustrated
here (Fig. 7B–D) for the irst time.
New CI record.
Leptodius planus Ward, 1934
(Figs. 4A, 7A)
Macromedaeus quinquedentatus (Krauss, 1843)
Leptodius planus Ward, 1934: 14, pl. 3 ig. 6, 6a
Xantho gracilis, Tweedie, 1950: 114
Leptodius gracilis, Balss, 1938: 52; Tweedie, 1947: 28; Morgan,
2000: 121 (table)
Material examined. Christmas I.: 1 ovig. ♀, 13.3 × 8.9
mm (ZRC), stn CI3-23.
Remarks. Macromedaeus quinquedentatus (type locality:
South Africa) has been reported from localities in the
Indian and Paciic oceans (see Serène, 1984). The key by
Serène (1984) separates this species from the similar M.
crassimanus by the proportions of the carapace width and
the fronto-orbital width, the dorsal areolation of the carapace,
and most indicatively, the G1 morphology. In the absence
of a male specimen, it appears that the best way to separate
M. quinquedentatus from M. crassimanus is by taking note
of the degree of areolation of the carapace regions. In M.
quinquedentatus, the areolae are sharper and more acute, and
the grooves are wider and deeper. In addition, the superoexternal margin of the chelar palm has four prominent
tubercles (versus merely rugose and without tubercles in
M. crassimanus), and the external surface of the chelar
palm has two longitudinal, granulate ridges (versus smooth
and without ridges in M. crassimanus). Furthermore, the
ovigerous female reported here is much smaller at maturity
than the preceding species.
Material examined. Christmas I.: 1 ♀, 7.2 × 4.8 mm
(ZRC), stn CI3-13.
Cocos (Keeling) Is.: 1 ♂, 17.5 × 11.1 mm (ZRC), stn CK108; 1 ♀, 10.2 × 7.0 mm (ZRC), stn CK2-21.
Remarks. Leptodius planus was originally described by
Ward (1934) based on specimens from Christmas Island.
Balss (1938) synonymised L. planus under Leptodius gracilis
Dana, 1852 (type locality: Wake Island, northern Paciic),
and the two are indeed similar in having a relatively smooth
carapace with the dorsal regions poorly deined, and in
having four teeth on the anterolateral margin of the carapace.
Although Ng et al. (2008), listed L. planus as distinct from
L. gracilis, they did not provide a reason for their action.
Initial unpublished results from an ongoing revision of the
genus Leptodius by the irst and last authors, in collaboration
with Lee Sang-kyu (Korea), show that the two are distinct
species. There appears to be a consistent difference in the
morphology of the distal portions of the G1 in these two
species: in L. gracilis the apical lobe is in a continuous curve
New CI record.
284
RAFFLES BULLETIN OF ZOOLOGY 2014
Fig. 5. Live colouration. A, Atergatis dilatatus De Haan, 1835, 1 ♂, 137.0 × 81.7 mm (ZRC), stn CI3-D04; B, Atergatis latissimus (H.
Milne-Edwards, 1834), 1 ♀, 28.6 × 18.2 mm (ZRC), stn CI1-31; C, Zosimus actaeoides (A. Milne-Edwards, 1867), 1 ♂, 17.8 × 12.2 mm
(ZRC), stn CK2-18; D, Zosimus aeneus (Linnaeus, 1758), 1 ♂ (ZRC), stn CK2-17; E, Zozymodes xanthoides (Krauss, 1843), 1 ♂, 7.1 ×
4.6 mm (ZRC), stn CK2-09; F, Zosimus aeneus (Linnaeus, 1758), 1 ♂ (ZRC), stn CI3-25.
285
Mendoza et al.: Xanthidae from Christmas and Cocos Islands
Nanocassiope alcocki (Rathbun, 1902)
(Fig. 4C)
also different from that recorded for P. elegans. In the colour
plates of T. Sakai (e.g., Sakai, 1935: pl. 47 ig. 1; 1976: pl.
155 ig. 2), P. elegans is shown to have a uniformly purplish
colouration in the dorsal aspect of carapace and pereopods,
except for the cheliped ingers which are dark brown. On
the other hand, the CI specimen is vividly reddish-orange,
with white and yellow mottling, and the ambulatory legs
have a whitish banding pattern at the joints between the
articles (Fig. 4D). This male has not been described as a
new species because examination of more specimens from
Christmas I. and also from other localities might provide
additional information. The irst author has observed similar
forms from the Philippines and Taiwan, which warrants a
comprehensive approach in elucidating the taxonomy of
this species.
Nanocassiope alcocki, Morgan, 2000: 121 (table).
Material examined. Christmas I.: 1 juv. ♂, 4.0 × 2.8
mm (ZRC), stn CI3-D02; 1 ♀, 9.8 × 6.5 mm (ZRC), stn
CI3-D08(157).
Remarks. This species was previously described from
the Maldives (Nallandu) by Rathbun (1902) and has been
recorded from Christmas Island by Morgan (2000), but
was neither illustrated nor remarked upon. The present
specimens agree well with the description and illustrations
for N. alcocki (cf. Rathbun, 1902: 128, igs. 9, 10; Serène,
1984: 209, pl. 28 ig. F). It is notable that this species was
collected at diving depth (<30 m) in Christmas Island, as it
is usually collected by trawl at greater depths (≥100 m) in
other localities (e.g., Serène, 1984: 209).
New CI record.
Xanthias cherbonnieri Guinot, 1964
(Figs. 4E, 7F)
Nanocassiope tridentata Davie, 1995
(Fig. 6B)
Material examined. Christmas I.: 1 ♂, 4.5 × 3.0 mm
(ZRC), stn CI2-D18.
Material examined. Cocos (Keeling) Is.: 1 ovig. ♀, 5.2 ×
3.5 mm (ZRC), stn CK2-12.
Remarks. This species was irst described from Aldabra,
and has been subsequently reported from Réunion (Guinot,
1964; Serène, 1984). The CI specimen agrees with the
description for X. cherbonnieri, particularly of the carapace
and ambulatory legs (Guinot, 1964: 34, pl. 2 figs. 1–4;
Serène, 1984: 196, pl. 27 ig. D). It differs from the type,
however, in having aberrant chelipeds where the palm
and ingers are much more slender than normal (Fig. 4E).
Furthermore, the abdomen appears wider than what is usually
seen in male Xanthias spp., and, although there is a pair of
well-developed G1s and G2s, the 3rd to 5th pleomeres also
have unspecialised, immature pleopods. The G1 (Fig. 7F)
is quite short and simple, with a few spiniform subterminal
setae. This G1 morphology is different from that of the type
species of the genus, X. granosomanus Dana, 1852 (= X.
lamarcki (H. Milne-Edwards, 1834)), where the G1 is quite
long, slender and has long, plumose subterminal setae, and
is, instead, more similar to that of X. latifrons (De Man,
1887) (cf. Serène, 1984: igs. 112, 117).
Remarks. This species was previously known only from
the type locality, Ambon Bay, Indonesia. In the present
specimen, a small, ovigerous female, the irst anterolateral
tooth after the exorbital tooth is much reduced, making the
carapace anterolateral margin appear tridentate. This is one
of the main diagnostic features of N. tridentata (cf. Davie,
1995: 205, ig. 2A).
New CK record.
Paraxanthias aff. elegans
(Figs. 4D, 7G)
Material examined. Christmas I.: 1 ♂, 7.5 × 5.1 mm
(ZRC), stn CI3-D02.
Remarks. The present specimen is most similar to
Paraxanthias elegans (Stimpson, 1858) (type locality:
Shimoda, Japan) in the subhexagonal carapace, the dorsal
surface of which is smooth and glabrous, with the regions
slightly defined. The carapace anterolateral margin is
likewise armed with four small, but well defined, teeth
posterior to the exorbital angle. The carpi of the chelipeds
are prominently granular (cf. Stimpson, 1907: pl. 5 ig. 3;
Sakai, 1976: ig. 226, pl.155 ig. 2; Dai & Yang, 1991: pl.
38 ig. 5). The G1 is slender and curved, with long, supple,
and plumose subterminal setae; the distal tip is bent at an
angle (cf. Forest & Guinot, 1961: ig. 72a, b; Dai & Yang,
1991: ig. 156B(3)).
New CI record.
Subfamily Zosiminae Alcock, 1898
Atergatis dilatatus De Haan, 1835
(Fig. 5A, 8A-C)
Material examined. Christmas I.: 1 ♂, 137.0 × 81.7 mm
(ZRC), stn CI3-D04.
Remarks. The present specimen, a large male, agrees well
with the description and illustrations of Atergatis dilatatus
De Haan, 1835 (type locality: seas of China or Japan, no
specific locality) (cf. De Haan, 1835: 46, pl. 14 fig. 2;
Serène, 1984: 148, pl. 21 ig. C). The G1 is illustrated here
for the irst time (Fig. 8A–C). This species has also been
The specimen from Christmas I. differs, however, in having
the external surface of the chelar palm smooth and without
longitudinal rows of granules, and in having relatively longer
and more slender ambulatory legs. The live colouration is
286
RAFFLES BULLETIN OF ZOOLOGY 2014
Fig. 6. Carapace, dorsal view (A, B). A, Etisus frontalis Dana, 1852, 1 ♀, 8.2 × 5.8 mm (ZRC), CK1-18; B, Nanocassiope tridentata
Davie, 1995, 1 ovig. ♀, 5.2 × 3.5 mm (ZRC), CK2-12. Left G1 (C–I). C, D, Cymo cerasma Morgan, 1990: 1 ♂, 10.6 × 9.6 mm (ZRC), stn
CI1-D04; E, Etisus demani Odhner, 1925: 1 ♂, 28.5 × 18.9 (ZRC) stn CI3-16; F, Etisus odhneri Takeda, 1971: 1 ♂, 12.5 × 8.8 mm (ZRC),
stn CK2-03; G–I, Liomera virgata (Rathbun, 1906): 1 ♂, 14.6 × 8.2 mm (ZRC), stn CI3-D02. Scale bars: A, B = 1.0 mm, C–I = 0.5 mm.
287
Mendoza et al.: Xanthidae from Christmas and Cocos Islands
Fig. 7. Left G1. A, Leptodius planus Ward, 1934: 1 ♂, 17.5 × 11.1 mm (ZRC), stn CK1-08. B–D, Lachnopodus ponapensis (Rathbun,
1907): 1 ♂, 18.6 × 13.0 mm (ZRC), stn CI3-23. E, Medaeus elegans A. Milne-Edwards, 1867: 1 ♂, 13.0 × 8.9 mm (ZRC), stn CK2-12.
F, Xanthias cherbonnieri Guinot, 1964: 1 ♂, 4.5 × 3.0 mm (ZRC), stn CI2-D18. G, Paraxanthias aff. elegans: 1 ♂, 7.5 × 5.1 mm (ZRC),
stn CI3-D02. Scale bars: A–G = 0.5 mm.
288
RAFFLES BULLETIN OF ZOOLOGY 2014
Fig. 8. Left G1. A–C, Atergatis dilatatus De Haan, 1835: 1 ♂, 137.0 × 81.7 mm (ZRC), stn CI3-D04. D, E, Zozymodes xanthoides (Krauss,
1843): 1 ♂, 7.1 × 4.6 mm (ZRC), stn CK2-09. F, G, Zosimus actaeoides (A. Milne-Edwards, 1867): 1 ♂, 19.7 × 13.4 mm (ZRC), stn
CI2-D18. Scale bars: A= 5.0 mm; B, C, F = 1.0 mm; D, E, G = 0.5 mm.
289
Mendoza et al.: Xanthidae from Christmas and Cocos Islands
Platypodia anaglypta (Heller, 1861)
(Fig. 4F)
reported from Japan, China, New Caledonia, Sri Lanka, the
Andamans, and Kenya (Serène, 1984).
Material examined. Christmas I.: 1 ♂, 13.6 × 9.0 mm
(ZRC), stn CI3-15(050).
New CI record.
Atergatis granulatus De Man, 1889
Remarks. This species is widespread in the Indo-West
Paciic region (Serène, 1984).
Material examined. Christmas I.: 1 carapace, 54.6 × 34.1
mm (ZRC), stn CI2-D05; 1 broken carapace (not measured)
(ZRC), stn CI3-D02(125).
New CI record.
Zosimus actaeoides (A. Milne-Edwards, 1867)
(Figs. 5C, 8F, G)
Remarks. This species was originally described from
Mauritius (De Man, 1889), and has also been reported from
the Red Sea and Japan (Serène, 1984). The entirely granular
carapace, with non-cristate anterolateral margins and no
epibranchial teeth, is diagnostic for this species. Only two
carapaces and no live specimens were collected, although the
good condition of one of the carapaces suggests that there
are probably live individuals occurring on Christmas Island.
A similar situation was observed for Neoliomera cerasinus
Ng, 2002 (type locality: Christmas Island), where congregates
of numerous elements of its exoskeleton (e.g. carapaces,
chelae, ambulatory legs) were found on the loor of certain
areas within the underwater caves around Christmas Island
(H.H. Tan, pers. comm.).
Zosimus actaeoides, Morgan, 2000: 121 (table)
Material examined. Christmas I.: 1 ♂, 19.7 × 13.4 mm
(ZRC), stn CI2-D18.
Cocos (Keeling) Is.: 1 ♂, 17.8 × 12.2 mm (ZRC), stn CK2-18.
Remarks. This species was first described from New
Caledonia (A. Milne-Edwards, 1867), and has also been
reported from the Hawaiian Islands (Rathbun, 1906). It was
also recorded by Morgan (2000) from Christmas Island.
One specimen each was collected from Christmas and the
Cocos (Keeling) islands, during the recent expeditions. The
present specimens generally agree with the description and
illustrations for Z. actaeoides (cf. A. Milne-Edwards, 1867:
273; 1873: pl. 7 ig. 7). The G1 is illustrated here for the
irst time (Fig. 8F, G).
New CI record.
Atergatis latissimus (H. Milne-Edwards, 1834)
(Fig. 5B)
New CK record.
Atergatis latissimus, Tweedie, 1947: 30; Morgan, 2000: 121 (table);
Davie, 2002: 562.
Zosimus aeneus (Linnaeus, 1758)
(Fig. 5D, F)
Material examined. Christmas I.: 2 ♀, 28.6 × 18.2 mm,
50.8 × 31.6 mm (ZRC), stn CI1-31.
Zozymus aeneus, Calman, 1909: 704; Tweedie, 1947: 27
Zoozymus aeneus, Tweedie, 1950: 115, ig. 1c
Zosimus aeneus, Morgan, 2000: 121 (table); Davie, 2002: 567
Remarks. This uncommon species was originally described
from Australia (H. Milne-Edwards, 1834), and has been
reported from Japan, the Marshall Islands, and Mauritius
(cf. Serène, 1984). Tweedie (1947: 30) recorded a large
male (CW = 122 mm) from Christmas Island, which, he
remarked, agreed well with the illustration of Atergatis
latissimus frontalis (De Haan, 1837) by T. Sakai (1939: pl.
88 ig. 1). He did not provide a igure, however. Morgan
(2000) also listed this species in his report, presumably based
on the record of Tweedie (1947). Following Tweedie (1947),
Atergatis frontalis (De Haan, 1837), is considered a junior
subjective synonym of A. latissimus (H. Milne-Edwards,
1834) (cf. Ng et al., 2008). The present specimens, both
female, are much smaller than Tweedie's material. The smaller
specimen still has the thin, whitish band on the anterolateral
and frontal margins of the carapace typical of the juvenile
form, igured here (Fig. 5B).
Material examined. Christmas I.: 1 ♂, 45.5 × 31.6 mm
(ZRC), 1 ♂, 50.6 × 33.4 mm (ZRC), stn CI1-13.
Cocos (Keeling) Is.: 2 ♀, 70.1 × 45.5 mm, 82.0 × 53.8 mm
(ZRC), stn CK1-02; 1 ♀, 82.5 × 56.1 mm (ZRC) stn CK1-08.
Remarks. This species is found throughout the IndoWest Paciic region, including the Christmas and Cocos
(Keeling) islands. The variation in live colouration is noted
here. Tweedie (1950: 115) noted the colour pattern of the
specimens found in the Cocos (Keeling) Is. as being “white
with the carapace and dorsal surface of the limbs marked
with a complex pattern of dark umber or very dark sienna
blotches; occasionally the white ground is suffused with pale
purple.” The same pattern was observed in the present Cocos
(Keeling) specimens, referred to in Table 1 as “bi-coloured”
(Fig. 5D). This is quite distinct from the normal colouration
of this species (as seen in the specimens from Christmas
I.), which usually shows more brown and purple and less
of the white background (Fig. 5F). Besides the differences
in colouration, no other morphological differences can be
discerned in the present specimens.
290
RAFFLES BULLETIN OF ZOOLOGY 2014
Zozymodes xanthoides (Krauss, 1843)
(Figs. 5E, 8D, E)
Borradaile LA (1902) Marine crustaceans. III. The Xanthidae
and some other crabs. In: St. Gardiner J (ed.) The Fauna and
Geography of the Maldive and Laccadive Archipelagoes, 1(3):
237–271.
Buitendijk AM (1941) Biological results of the Snellius Expedition.
XIII. On some Xanthidae, chiely of the genus Platypodia Bell.
Temminckia, 6: 295–312.
Calman WT (1909) On decapod Crustacea from Christmas Island,
collected by Dr. C.W. Andrews, F.R.S., F.Z.S. Proceedings
of the Zoological Society of London, 1909: 703–713, pl. 72.
Clark PF & Galil BS (1993) A revision of the xanthid genus Pilodius
Dana, 1851 (Crustacea: Brachyura: Xanthoidea). Journal of
Natural History, 27: 1119–1206.
Dai A & Yang S (1991) Crabs of the China Seas. China Ocean
Press, Beijing; Springer-Verlag, Berlin, Heidelberg, New York,
Tokyo, 682 pp., 74 pls.
Dai AY, Yang SL, Song YZ & Chen GX (1986) Crabs of the China
Seas. China Ocean Press, Beijing, pp. 1–11, 1–642. [in Chinese]
Dana JD (1851) On the classiication of the Cancroidea. American
Journal of Science and Arts, 12(34): 121–131.
Dana JD (1852) Crustacea. Part I. United States Exploring
Expedition. During the Years 1838, 1839, 1840, 1841, 1842.
Under the Command of Charles Wilkes, U.S.N. Vol. 13. C.
Sherman, Philadelphia, 685 pp.
Davie PJF (1995) Two new species of Nanocassiope from the
Western Paciic (Crustacea, Brachyura, Xanthidae). Bulletin du
Muséum national d’Histoire naturelle, Paris, 4e sér., 17: 201–210.
Davie PJF (1997) Crustacea Decapoda: deep water Xanthoidea
from the south-western Paciic and the western Indian Ocean.
In: Crosnier A (ed.) Resultats des campagnes MUSORSTOM,
Volume 18. Memoires du Muséum national d’Histoire naturelle,
176: 337–387.
Davie PJF (2002) Crustacea: Malacostraca: Eucarida (Part 2):
Decapoda – Anomura, Brachyura. In: Wells A & Houston
WWK (eds.) Zoological Catalogue of Australia, Vol. 19.3B.
CSIRO Publishing, Melbourne, xiv + 641 pp.
Edmondson CH (1962) Xanthidae of Hawaii. Occasional Papers
of the Bernice P. Bishop Museum, 20(13): 183–243.
Forest J & Guinot D (1961) Crustacés Décapodes Brachyoures
de Tahiti et des Tumaotu. In: Expédition Française sur les
Récifs Coralliens de la Nouvelle Calédonie. Muséum national
d’Histoire naturelle, Paris, 195 pp., 18 pls.
Galil B & Vannini M (1990) Research on the coast of Somalia.
Xanthidae, Trapeziidae, Capiliidae, Menippidae (Crustacea
Brachyura). Tropical Zoology, 3: 21–56.
Garth JS, Haig J & Knudsen JW (1987) Crustacea Decapoda
(Brachyura and Anomura) of Eniwetak Atoll. In: Devaney DM,
Reese ES, Burch BL & Helfrich P (eds.) The Natural History
of Eniwetak Atoll. Vol. 2. Biogeography and Systematics.
U.S. Department of Energy, Ofice of Scientiic and Technical
Information, Oak Ridge, Tennesse, pp. 235–261.
George RW (1978) The land and freshwater crabs of Christmas
Island. Western Australian Museum, Perth, 22 pp.
Guinot D (1964) Crustacés décapodes brachyoures (Xanthidae)
des campagnes de la Calypso en Mer Rouge (1952), dans
le Golfe Persique et a l’Ile Aldabra (1954). Mémoires du
Muséum national d’Histoire naturelle, sér. A, 32(1): 1–108,
index, pls. 1–12.
Guinot D (1967) Recherches préliminaires sur les groupements
naturels chez les crustaces décapodes brachyoures. II. Les
anciens genres Micropanope Stimpson et Medaeus Dana.
Bulletin du Muséum national d’Histoire naturelle, 2e sér., 39:
345–374.
Guinot D (1969) Sur divers Xanthidae, notamment sur Actaea de
Haan et Paractaea gen. nov. (Crustacea Decapoda Brachyura).
Cahiers du Paciique, 13: 223–267.
Material examined. Cocos (Keeling) Is.: 1 ♀, 8.6 × 5.5 mm
(ZRC), stn CK1-02; 1 ♂, 7.1 × 4.6 mm (ZRC), stn CK2-09.
Remarks. This species was irst described by Krauss (1843)
from Natal, South Africa. It has also been reported from
Madagascar, the Red Sea, the eastern coast of Africa from
Somalia to South Africa, and French Polynesia (Serène,
1984; Galil & Vannini, 1990; Poupin, 2010). The present
specimens agree well with the description and illustrations
in Serène (1984). The fresh colouration of this species is
recorded here for the irst time (Fig. 4H).
New CK record.
ACKNOWLEDGEMENTS
The authors are grateful for the support from the Christmas
Island National Park, the Pulu Keeling National Park, the
Queensland Museum and the Rafles Museum of Biodiversity
Research during the 2010–2012 expeditions to Christmas
Island and the Cocos (Keeling) Islands. We also thank
the expedition team members who were instrumental in
collections and data gathering on-site: Tan Heok Hui, Tan
Swee Hee, Joelle Lai, Tan Kai-xin, Tan Siong Kiat (NUS,
Singapore); Peter Davie (Queensland Museum, Brisbane);
Tohru Naruse, Yoshihisa Fujita (University of the Ryukyus,
Okinawa), Shih Hsi-Te (National Chung Hsing University,
Taiwan) and Max Orchard (CI National Park). We thank Tan
Heok Hui and Tohru Naruse for providing some of the colour
photographs used in this paper, Martyn Low for help with
literature search, and Lee Bee Yan for assistance with some
specimen measurements. Finally we thank Shane Ahyong,
Paul Clark & Tohru Naruse for their valuable comments on
the drafts of this paper.
LITERATURE CITED
Adams A & White A (1849) Crustacea. Part 2. In: Adams A (ed.)
The Zoology of the Voyage of H.M.S. Samarang Under the
Command of Captain Sir Edward Belcher, During the Years
1843–1846. Benham & Reeve, London, pp. i–viii, 33–67,
pls. 7–13.
Alcock A (1898) Materials for a carcinological fauna of India. No.
3. The Brachyura Cyclometopa. Part I. The family Xanthidae.
Journal of the Asiatic Society of Bengal, Calcutta, 67(2): 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 des caractères naturels
des genres avec la distinction des espèces. Description de
l’Egypte ou recueil des observations et des recherches qui ont
été faites en Egypte pendant l’expédition de l’armée française.
Histoire naturelle, Paris, 1(4): 77–98.
Balss H (1934) Die krabben der reise J.W. Harms’ nach der
Christmas-Insel und dem Malaiischen Archipel. Zoologischer
Anzeiger, 106(10): 225–237.
Balss H (1938) Ueber einige Xanthidae (Crustacea, Dekapoda) von
Singapore und Umgebung. Bulletin of the Rafles Museum,
14: 48–63, pls. 2, 3.
291
Mendoza et al.: Xanthidae from Christmas and Cocos Islands
Mendoza JCE & Ng PKL (2011) The Polydectinae Dana, 1851,
of the Philippines, with description of a new genus for Lybia
hatagumoana Sakai, 1961 (Crustacea: Decapoda: Brachyura:
Xanthidae). Zootaxa, 3052: 51–61.
Miers EJ (1884) Crustacea. Report on the zoological collections
made in the Indo-Paciic Ocean during the voyage of H.M.S.
‘Alert’ 1881–1882. Part I. The collections from Melanesia.
Part II. The collections from the Western Indian Ocean. British
Museum, London, pp. 178–322, 513–575, pls. 18–32, 46–51.
Milne-Edwards A (1865) Etudes zoologiques sur les Crustacés
récents de la famille des Cancériens. Nouvelle Archives du
Muséum d’Histoire naturelle, Paris, 1: 177–308, pls. 11–19.
Milne-Edwards A (1867) Descriptions de quelques espèces nouvelles
de Crustacés Brachyures. Annales de la Societé Entomologique
de France, 4e sér., 7: 263–288.
Milne-Edwards A (1873) Recherches sur la faune carcinologique
de la Nouvelle-Calédonie, Deuxième Partie. Nouvelles Archives
du Muséum d’Histoire naturelle, Paris, 9: 155–332, pls. 4–18.
Milne-Edwards H (1834) Histoire naturelle des Crustacés,
comprenant l’anatomie, la physiologie et la classiication de
ces animaux. Libraire Encyclopédique de Roret. Vol. 1. Roret,
Paris, 468 pp.
Morgan GJ (1990) A collection of Thalassinidea, Anomura and
Brachyura (Crustacea: Decapoda) from the Kimberley Region of
northwestern Australia. Zoologische Verhandelingen, 265: 1–90.
Morgan GJ (2000) Decapod Crustacea of Christmas Island. In:
Berry PF & Wells FE (eds.) Survey of the Marine Fauna of the
Montebello Islands, Western Australia and Christmas Island,
Indian Ocean. Records of the Western Australian Museum,
Supplement 59: 117–123.
Ng PKL (2002) On a new species of cavernicolous Neoliomera
(Crustacea: Decapoda: Brachyura: Xanthidae) from Christmas
Island and Ryukyus, Japan. Rafles Bulletin of Zoology, 50(1):
95–99.
Ng PKL & Chia DGB (1994) The genus Glyptocarcinus Takeda,
1973, with descriptions of a new subfamily, two new genera and
two new species from New Caledonia (Crustacea: Decapoda:
Brachyura: Xanthidae). Rafles Bulletin of Zoology, 42(3):
701–731.
Ng PKL & Clark PF (2002) Description of a new species of
Paramedaeus Guinot, 1967, with notes on Paramedaeus
simplex (A. Milne-Edwards, 1873) and Metaxanthops acutus
Serène, 1984 (Decapoda, Brachyura, Xanthoidea, Xanthidae).
Crustaceana, 75(3–4): 527–538.
Ng PKL, Guinot D & Davie PJF (2008) Systema Brachyurorum:
Part I. An annotated checklist of extant brachyuran crabs of
the world. Rafles Bulletin of Zoology, Supplement 17: 1–286.
Ng PKL & Holthuis LB (2007) Etisus H. Milne-Edwards, 1834 and
Chlorodiella Rathbun, 1897 (Crustacea, Decapoda, Brachyura):
proposed conservation of the generic names by suppression of
the generic name Clorodius A.G. Desmarest, 1823. Bulletin of
Zoological Nomenclature, 64(1): 19–24.
Odhner T (1925) Monographierte Gattungen der Krabben-familie
Xanthidae. I. Göteborgs Kungliga Vetenskaps- och VitterhetsSamhälles Handlingar, (4)29(1): 1–92, igs. 1–7, pls. 1–5.
Orchard M (2012) Crabs of Christmas Island. Christmas Island
Natural History Association, 288 pp.
Ortmann A (1893) Die Decapoden-Krebse des Strassburger
Museums, mit besonderer Berücksichtigung der von Herrn Dr.
Döderlein bei Japan und bei den Liu-Kiu-Inseln gesammelten
und zur Zeit im Strassburger Museum aufbewahrten. VII.
Theil. Abtheilung: Brachyura (Brachyura genuina Boas) II.
Unterabtheilung: Cancroidea, 2. Section: Cancrinea, 1. Gruppe:
Cyclometopa. Zoologische Jahrbücher Abteilung für Systematik,
Geographie und Biologie der Thiere, 7: 411–495, pl. 17.
Guinot D (1976) Constitution de quelques groups naturels chez
les crustacés décapodes brachyoures. I. La superfamille des
Bellioidea et trois sous-familles de Xanthidae (Polydectinae
Dana, Trichiinae de Haan, Actaeinae Alcock). Mémoires du
Muséum national d’Histoire naturelle, sér. A, 97: 1–308, pls.
1–19.
Haan W de (1833–1849) Crustacea. In: von Siebold PF (ed.)
Fauna Japonica, sive Descriptio animalium, quae in itinere per
Japoniam, jussu et auspiciis superirum, qui summum in India
Batava imperium tenent, suscepto, annis 1823–1830 collegit,
notis, observationibus et adumbrationibus illustravit P. F. de
Siebold. Conjunctis studiis C. J. Temminck et H. Schlegel pro
Vertebratis atque W. de Haan pro Invertebratis elaborata Regis
auspicis edita. Leiden, Lugundi-Batavorum: P.F. v. Siebold.
Decas II: 25–64, pls. 9–15, 17, C, D.
Heller C (1861) Synopsis der im rothen Meere vorkommenden
Crustaceen. Verhandlungen der Zoologisch-Botanischen
Gessellschaft in Wien, 11: 3–32.
Herbst JFW (1785) Versuch einer Naturgeschichte der Krabben
und Krebse, nebst einer systematischen Beschreibung ihrer
verschiedenen Arten. Joh. Casper Feussly, Zürich, 1(6):
183–206, pls. 10–13.
Herbst JFW (1790) Versuch einer Naturgeschichte der Krabben
und Krebse, nebst einer systematischen Beschreibung ihrer
Arten. Gottlieb, August & Lange, Berlin & Stralsund, 1(8):
239–274, pls. 18–21.
Krauss F (1843) Die Südafrokanischen Crustaceen. Ein
Zusammenstellung aller bekannten Malacostraca Bemerkungen
über deren Lebensweise und geographische Verbreitung, nebst
Beschreibung und Abbildung mehrerer neuen Arten. Stuttgart,
68 pp., pls. 1–4.
Lasley RM Jr & Ng PKL (2013) A new species of the crab genus
Zozymodes (Crustacea: Decapoda: Xanthidae) from Guam.
Micronesica, 2013-03: 1–12.
Linnaeus C (1758) Systema Naturae per Regna Tria Naturae,
Secundum Classes, Ordines, Genera, Species, cum Characteribus,
Differentiis, Synonymis, Locis. Tom. 1. Laurentii Salvii,
Holmiae,iii + 824 pp.
MacLeay WS (1838) Illustrations of the Annulosa of South Africa;
being a portion of the objects of natural history chiely collected
during an expedition into the interior of South Africa, under
the direction of Dr. Andrew Smith, in the years 1834, 1835,
and 1836; itted out by the “Cape of Good Hope Association
for Exploring Central Africa.” In: Smith A (ed.) Illustrations
of the Zoology of South Africa investigations. Smith, Elder
and Co., London, pp. 1–75, pls. 1–4.
Man JG de (1887) Report on the podophthalmous Crustacea of the
Mergui Archipelago, collected for the Trustees of the Indian
Museum, Calcutta, by Dr. John Anderson, F.R.S., superintendent
of the museum. Parts I–V. Journal of the Linnean Society of
London (Zoology), 22(136–137): 1–128.
Man JG de (1889) Über einige neue oder seltene indopaciische
Brachyuren. Zoologische Jarbücher. Abteilung für Systematik,
Geographie und Biologie der Thiere, 4: 409–452, pls. 9, 10.
Mendoza JCE & Guinot D (2011) Revision of the genus
Glyptoxanthus A. Milne-Edwards, 1879, and establishment
of Glyptoxanthinae nov. subfam. (Crustacea: Decapoda:
Brachyura: Xanthidae). Zootaxa, 3015: 29–51.
Mendoza JCE & Manuel-Santos MR (2012) Revision of Garthiella
Titgen, 1986 (Crustacea: Decapoda: Brachyura: Xanthidae),
with description of a new subfamily and a new species from
the central Philippines. Zootaxa, 3446: 32–48.
Mendoza JCE & Ng PKL (2010) The euxanthine crabs (Crustacea:
Brachyura: Xanthidae) of the Philippines. Rafles Bulletin of
Zoology, 58(1): 57–74.
292
RAFFLES BULLETIN OF ZOOLOGY 2014
Poupin J (2010) Biodiversité de l’Indo-Paciique tropical français:
2514 espèces de crustacés décapodes et stomatopodes. Rapport
scientiique de l’Institut de Recherche de l’Ecole Navale, 76 pp.
Rathbun MJ (1894) Descriptions of two new species of crabs from
the Western Indian Ocean, presented to the National Museum
by Dr. W. L. Abbott. Proceedings of the United States National
Museum, 17(979): 21–24.
Rathbun MJ (1897) A revision of the nomenclature of the Brachyura.
Proceedings of the Biological Society of Washington, 11:
153–167.
Rathbun MJ (1902) Crabs from the Maldive Islands. Bulletin of
the Museum of Comparative Zoology, Harvard College, 39(5):
123–137, 1 pl.
Rathbun MJ (1906) The Brachyura and Macrura of the Hawaiian
Islands. Bulletin of the United States Fish Commission, 23(3):
827–930, pls. 1–24.
Rathbun MJ (1907) Reports on the scientiic results of the expedition
to the tropical Paciic, in charge of Alexander Agassiz, by the
U.S. Fish Commission steamer “Albatross”, from August,
1899, to March, 1900, Commander Jefferson F. Moser,
U.S.N., commanding. IX. Reports on the scientiic results of
the expedition to the tropical Paciic, in charge of Alexander
Agassiz, by the U.S. Fish Commission steamer “Albatross”,
from October, 1904, to March, 1905, Lieut.-Commander L.M.
Garrett, U.S.N., commanding. X: The Brachyura. Memoirs
of the Museum of Comparative Zoology at Harvard College,
35(2): 25–74.
Rathbun MJ (1911) Marine Brachyura. In: The Percy Sladen Trust
Expedition to the Indian Ocean in 1905 under the leadership
of Mr. J. Stanley Gardiner. Vol. III. No. IX. Transactions of
the Linnaean Society of London, Zoology, 14(2): 191–261,
pls. 15–20.
Sakai K (1999) J.F.W. Herbst – collection of decapod Crustacea
of the Berlin Zoological Museum, with remarks on certain
species. Naturalists, Tokushima Biological Laboratory, Shikoku
University, 6: 1–45, pls. 1–21.
Sakai T (1935) New or rare species of Brachyura, collected by the
Misago during the zoological survey around the Izu Peninsula.
Science Reports of the Tokyo Bunrika Daigaku, 2(32): 63–88,
pls. 6–8.
Sakai T (1939) Studies on the crabs of Japan. IV. Brachygnatha,
Brachyrhyncha. Yokendo Co., Tokyo, pp. 365–741, pls. 42–111.
Sakai T (1976) Crabs of Japan and the Adjacent Seas. Kodansha
Ltd, Tokyo. [In 3 volumes: 1, English text: i–xxix, 1–773, igs.
1–379; 2, Plates volume: 1–16, pls. 1–251; 3, Japanese text:
1–461, igs. 1–2, 3 maps]
Serène R (1972) Observations on the Indo-pacific species of
Kraussia Dana, 1852 (Decapoda: Brachyura). Journal of the
Royal Society of Western Australia, 55(2): 41–62.
Serène R (1984) Crustacés Décapodes Brachyoures de l’Ocean
Indien occidental et de la Mer Rouge. Xanthoidea: Xanthidae
et Trapeziidae. Avec un Addendum par Crosnier, A. Carpiliidae
et Menippidae. Faune tropicale, 24: 1–349, pls. 1–48.
Stimpson W (1858) Prodromus descriptionis animalium
evertebratorum, quae in Expeditione ad Oceanum Paciicum
Septentrionalem, a Republica Federata missa, Cadwaladaro
Ringgold et Johanne Rodgers Ducibus, observavit et descripsit.
Pars IV. Crustacea Cancroidea et Corystoidea. Proceedings of
the Academy of Natural Sciences of Philadelphia, 10: 31–40.
Stimpson W (1860) Notes on North American Crustacea in the
Museum of the Smithsonian Institution. No. II. Annals of
the Lyceum of Natural History of New York, 7: 177–246
[49–118], pls. 2, 5.
Stimpson W (1907) Report on the Crustacea (Brachyura and
Anomura) collected by the North Paciic Exploring Expedition,
1853–1856. Smithsonian Miscellaneous Collections, 49: 1–240,
26 pls.
Takeda M (1971) New and rare crabs from the Palau Islands.
Micronesica, 7(1–2): 185–213.
Tan HH, Tan SK, Tan K, Lai JCY, Mendoza JC & Tan SH (2014)
Christmas Island and Pulu Keeling Expeditions 2010 to 2012.
Field work and locality records. Rafles Bulletin of Zoology,
Supplement 30: this issue.
Tweedie MWF (1947) On the Brachyura of Christmas Island.
Bulletin of the Rafles Museum, 18: 27–42.
Tweedie MWF (1950) The fauna of Cocos-Keeling Islands,
Brachyura and Stomatopoda. Bulletin of the Rafles Museum,
22: 105–148, pls. 16–17.
Ward M (1934) Notes on a collection of crabs from Christmas
Island, Indian Ocean. Bulletin of the Raffles Museum, 9:
5–28, pls. 1–3.
Ward M (1941) New Brachyura from the Gulf of Davao, Mindanao,
Philippine Islands. American Museum Novitates, 1104: 1–15.
White A (1848) Short descriptions of new or little-known decapod
Crustacea. Proceedings of the Zoological Society of London,
1847: 222–228.
Wood Jones F (1909) The fauna of the Cocos-Keeling Atoll,
collected by F. Wood Jones. Proceedings of the Zoological
Society of London, 1909: 132–160.
Zehntner L (1894) Voyage de MM. M. Bedot et C.Pictet dans
l’Archipel Malais. Crustacés de l’Archipel Malais. Revue
Suisse de Zoologie et Annales du Musée d’Histoire Naturelle
de Genève, 2: 135–214, pls. 7–9.
293
Mendoza et al.: Xanthidae from Christmas and Cocos Islands
Table 1. Updated checklist of xanthid species found in Christmas and Cocos (Keeling) islands, with references and including
station records for species collected by the recent expeditions (2010–2012). Newly recorded species are indicated in bold
and with superscripts to indicate locality, e.g., Christmas I. new record = *CI; Cocos (Keeling) Is. new record = *CK.
Species Current Name
(sensu Ng et al., 2008)
Christmas Island (CI)
Record
Recent CI Stations
(2010–2012)
Cocos (Keeling)
Islands (CK)
Record
Recent CK Stations
(2010–2012)
ACTAEINAE
Actaeodes consobrinus (A.
Milne-Edwards, 1873)
Ward, 1934 (as Actaea
suffuscula Rathbun); Tweedie,
1947; Morgan, 2000
Actaeodes cf. hirsutissimus
(Rüppell, 1830)
Morgan, 2000
Actaeodes tomentosus (H.
Milne-Edwards, 1834)*CK
Calman, 1909 (as Actaea
tomentosa); Ward, 1934 (as
Actaea tomentosa); Tweedie,
1947 (as Actaea tomentosa);
Morgan, 2000
Epiactaea nodulosa (White,
1848)
Morgan, 2000
CI3-17
CI1-03,
CI1-31,
CI2-17,
CI3-07,
CI3-16,
CI3-23,
Tweedie, 1950
CI1-13,
CI2-09,
CI2-22,
CI3-15,
CI3-17,
CI3-25
This paper
CK1-02
Gaillardiellus rueppellii
(Krauss, 1843)*CK
This paper
CK2-09
Gaillardiellus superciliaris
(Odhner, 1925)
Tweedie, 1950
(as Actaea
superciliaris)
CK1-08
Paractaea plumosa Guinot,
in Sakai, 1976*CI
This paper
Paractaea rufopunctata (H.
Milne-Edwards, 1834) sensu
lato
Calman, 1909 (as Actaea
rufopunctata); Ward, 1934
(as Actaea rufopunctata);
Tweedie, 1947 (as Actaea
rufopunctata); Morgan, 2000
Tweedie, 1950
(as Actaea
rufopunctata)
Paractaea rufopunctata
rufopunctata (H. MilneEdwards, 1834)
Davie, 2002
Davie, 2002
Psaumis cavipes (Dana, 1852)
Ward, 1934 (as Actaea
fossulata Girard); Morgan,
2000
CI3-15, CI3-16,
CI3-17, CI3-23
Wood Jones,
1909 (as Actaea
fossulata); Tweedie,
1950 (as Actaea
cavipes)
Pseudoliomera granosimana
(A. Milne-Edwards, 1865)
Ward, 1934 (as Pseudliomera
natalensis Ward); Tweedie,
1947; Morgan, 2000
CI2-17
Tweedie, 1950
Pseudoliomera lata
(Borradaile, 1902)
Balss, 1934 (as Actaea lata);
Morgan, 2000
CI3-17
Pseudoliomera speciosa
(Dana, 1852)
Calman, 1909 (as Actaea
speciosa); Balss, 1934 (as
Actaea speciosa); Tweedie,
1947 (as Actaea speciosa);
Morgan, 2000
CI2-D03
Pseudoliomera variolosa
(Borradaile, 1902)
Morgan, 2000
Pseudoliomera violacea (A.
Milne-Edwards, 1873)*CI
This paper
CI1-D02, CI2-D03,
CI2-D15
CI2-D17, CI3-D05
294
Tweedie, 1950 (as
Actaea speciosa)
CK2-21
RAFFLES BULLETIN OF ZOOLOGY 2014
Species Current Name
(sensu Ng et al., 2008)
Christmas Island (CI)
Record
Recent CI Stations
(2010–2012)
Cocos (Keeling)
Islands (CK)
Record
Recent CK Stations
(2010–2012)
CHLORODIELLINAE
Tweedie, 1950;
Davie, 2002
Chlorodiella barbata
(Borradaile, 1900)
Chlorodiella cytherea Dana,
1852*CI, CK
This paper
CI1-03,
CI2-09,
CI2-22,
CI3-14,
CI3-16,
CI3-23,
Chlorodiella laevissima
(Dana, 1852)
Calman, 1909 (as Chlorodius
laevissimus); Tweedie, 1947;
Morgan, 2000
CI2-22, CI3-15,
CI3-17
Chlorodiella nigra (Forskål,
1775)
Calman, 1909 (as Chlorodius
niger); Tweedie, 1947;
Morgan, 2000
Cyclodius drachi (Guinot,
1964)*CI
This paper
CI2-D03
Cyclodius nitidus (Dana,
1852)
Calman, 1909 (as Phymodius
sculptus A.M.E.); Ward,
1934 (as Phymodius sculptus
A.M.E.); Tweedie, 1947 (as
Phymodius nitidus); Morgan,
2000; Davie, 2002
CI3-15, CI3-16,
CI3-17
CI1-31,
CI2-17,
CI3-13,
CI3-15,
CI3-17,
CI3-25
This paper
CK1-08, CK2-21
Tweedie, 1950
CK2-07
Tweedie, 1950
(as Phymodius
monticulosus Dana)
Cyclodius obscurus (Hombron
& Jacquinot, 1846)
Cyclodius ungulatus (H.
Milne-Edwards, 1834)
Calman, 1909 (as Cyclodius
gracilis Dana); Tweedie,
1947 (as Cyclodius gracilis);
Morgan, 2000 (as Phymodius
ungulatus); Davie, 2002
CI3-14, C I3-15,
CI3-16, CI3-25
Liocarpilodes harmsi (Balss,
1934)
Balss, 1934 (as Pilodius
harmsi Balss); Ward, 1934
(as Chlorodopis natalensis);
Tweedie, 1947 (as Pilodius
harmsi); Morgan, 2000;
Davie, 2002
CI1-13,
CI1-33,
CI3-17,
CI3-26,
Liocarpilodes integerrimus
(Dana, 1852)
Morgan, 2000
CI3-17
Pilodius areolatus (H. MilneEdwards, 1834)
Calman, 1909 (as
Chlorodopsis areolata);
Tweedie, 1947 (as
Chlorodopsis areolata);
Morgan, 2000
CI1-D18/19, CI313, CI3-14, CI3-16,
CI3-23
Pilodius lavus Rathbun,
1894*CI
This paper
CI1-D17
Tweedie, 1950
(as Phymodius
ungulatus)
CK1-08, CK1-18,
CK1-22, CK2-19,
CK2-21
Tweedie, 1950
(as Chlorodopsis
areolata)
CK1-02, CK1-08,
CK2-17, CK2-19,
CK2-21
CI1-20,
CI3-15,
CI3-22,
CI3-30
Tweedie, 1950
(as Chlorodopsis
melanodactyla
A.M.E.); Davie,
2002
Pilodius pubescens Dana,
1852
295
Mendoza et al.: Xanthidae from Christmas and Cocos Islands
Recent CI Stations
(2010–2012)
Cocos (Keeling)
Islands (CK)
Record
Species Current Name
(sensu Ng et al., 2008)
Christmas Island (CI)
Record
Recent CK Stations
(2010–2012)
Pilodius scabriculus Dana,
1852
Calman, 1909 (as
Chlorodopsis venusta
Rathbun); Tweedie, 1947
(as Chlorodiella venusta);
Morgan, 2000; Davie, 2002
Pilodius aff. spinipes Heller,
1861
Morgan, 2000
Sulcodius cf. delexus (Dana,
1852)
Morgan, 2000 (as Etisus cf.
delexus)
Tweediea odhneri (Gordon,
1934)
Ward, 1934 (as Tweediea
noelensis Ward); Tweedie,
1947 (as Phymodius odhneri);
Morgan, 2000
CI1-D16, CI2-D15,
CI2-D17, CI2-D18,
CI3-17, CI3-D02
Cymo andreossyi (Audouin,
1826)*CI
This paper
CI3-16
Tweedie, 1950
Cymo cerasma Morgan,
1990*CI, CK
This paper
CI1-D04
This paper
CK1-08
Cymo melanodactylus Dana,
1852
Calman, 1909; Tweedie,
1947; Morgan, 2000; Davie,
2002
CI3-16
Tweedie, 1950
CK2-21
Cymo quadrilobatus Miers,
1884*CI
This paper
CI1-D14(126),
CI1-D17,
CI1-D18/19,
CI2-D14
Tweedie, 1950;
Davie, 2002
CK2-12
Etisus albus (Ward, 1934)
Ward, 1934 (as Etisodes albus
Ward); Morgan, 2000; Davie,
2002
CI2-22
Etisus anaglyptus H. MilneEdwards, 1834
Morgan, 2000
Etisus bifrontalis
(Edmondson, 1935)
Davie, 2002
Etisus demani Odhner,
1925*CK
Tweedie, 1947; Morgan,
2000; Davie, 2002
CI2-22, CI3-16
This paper
CK1-18
Etisus dentatus (Herbst,
1785)*CI
This paper
CI3-14, CI3-16
Tweedie, 1950;
Davie, 2002
CK2-21
Etisus frontalis Dana,
1852*CK
This paper
CK1-18
Etisus laevimanus Randall,
1840
Tweedie, 1950;
Davie, 2002
CI3-14, CI3-15,
CI3-16, CI3-17,
CI3-23
CYMOINAE
ETISINAE
Etisus odhneri Takeda,
1971*CI, CK
This paper
CI1-D17, CI2-D12
Etisus splendidus Rathbun,
1906*CI
This paper
CI1-D17
296
This paper
CK2-03
RAFFLES BULLETIN OF ZOOLOGY 2014
Species Current Name
(sensu Ng et al., 2008)
Christmas Island (CI)
Record
Recent CI Stations
(2010–2012)
Cocos (Keeling)
Islands (CK)
Record
Recent CK Stations
(2010–2012)
EUXANTHINAE
Danielea noelensis (Ward,
1934)
Ward, 1934 (as Medaeus
noelensis Ward); Tweedie,
1947 (as Medaeus granulosus
Haswell); Morgan, 2000;
Davie, 2002 (as Paramedaeus
noelensis)
Tweedie, 1950
(as Euxanthus
exsculptus Herbst);
Davie, 2002
(as Euxanthus
exsculptus Herbst)
Euxanthus aff. exsculptus
Medaeus elegans A. MilneEdwards, 1867*CI, CK
This paper
Miersiella haswelli (Miers,
1886)
Davie, 1997, 2002
CI3-D01
Paramedaeus octogesimus Ng
& Clark, 2002
Paramedaeus sp.*CI
This paper
CI3-D08
Ward, 1934 (as Kraussia
proporcellana Ward);
Tweedie, 1947; Morgan,
2000; Davie, 2002
CI1-31(184), CI323
This paper
CK2-12, CK2-13
Ng & Clark, 2002
CK1-18
Tweedie, 1950
CK2-05, CK2-17
KRAUSSIINAE
Kraussia rugulosa (Krauss,
1843)
Palapedia integra (De Haan,
1835)
Davie, 2002
Palapedia rastripes (Müller,
1887)
Tweedie, 1950 (as
Palapedia ?integra
(De Haan, 1835));
Serène, 1972;
Davie, 2002
LIOMERINAE
Bruciana cf. pediger
Morgan, 2000 (as Liomera cf.
pediger)
Liomera bella (Dana, 1852)
Calman, 1909 (as Carpilodes
vaillantianus A.M.E.); Balss,
1934 (as Carpilodes bella);
Ward, 1934 (as Carpilodes
bellus); Tweedie, 1947 (as
Carpilodes bellus); Morgan,
2000
CI1-13, CI1-31,
CI1-D18/19, CI209, CI2-17, CI2-22,
CI3-14, CI3-15,
CI3-16, CI3-23,
CI3-25
CK1-02, CK2-21
Tweedie, 1950
(as Carpilodes
caelatus); Davie,
2002
Liomera caelata (Odhner,
1925)
Liomera cinctimana (White,
1847)
Tweedie, 1950 (as
Carpilodes bellus)
Balss, 1934; Morgan, 2000
CI2-D17
Tweedie, 1950 (as
Carpilodes laevis);
Davie, 2002
Liomera laevis (A. MilneEdwards, 1873)
297
CK1-02, CK1-18,
CK1-19, CK2-05,
CK2-21
Mendoza et al.: Xanthidae from Christmas and Cocos Islands
Species Current Name
(sensu Ng et al., 2008)
Christmas Island (CI)
Record
Recent CI Stations
(2010–2012)
Liomera monticulosa (A.
Milne-Edwards, 1873)
Calman, 1909 (as Carpilodes
cariosus Alcock); Tweedie,
1947 (as Carpilodes
monticulosus); Morgan, 2000
CI2-D03, CI2-D06,
CI2-D10, CI3-D04
Liomera pallida (Borradaile,
1900)
Morgan, 2000
CI2-17, CI3-13,
CI3-25
Liomera rugata (H. MilneEdwards, 1834)
Calaman, 1909 (as Carpilodes
rugatus); Ward, 1934 (as
Carpilodes rugatus); Tweedie,
1947 (as Carpilodes rugatus);
Morgan, 2000; Davie, 2002
CI1-31, CI3-25
Liomera stimpsonii (A.
Milne-Edwards, 1865)*CI
This paper
CI2-D03, CI2-D15,
CI2-D18, CI2-D18,
CI3-D05
Recent CK Stations
(2010–2012)
Tweedie, 1950
(as Carpilodes
pallidus); Davie,
2002
Tweedie, 1950
(as Carpilodes
stimpsoni); Davie,
2002
Tweedie, 1950 (as
Carpilodes tristis);
Davie, 2002
Liomera tristis (Dana, 1852)
Liomera virgata (Rathbun,
1906)*CI
This paper
Liomera sp.
Morgan, 2000
Neoliomera cerasinus Ng,
2002
Ng, 2002
CI2-D17, CI3-D02
CI1-D06, CI1-D08,
CI2-D10, CI3-D02,
CI3-D04
Wood Jones,
1909 (as Liomera
pubescens)
Neoliomera pubescens (H.
Milne-Edwards, 1834)
Neoliomera richteroides
Sakai, 1969
Cocos (Keeling)
Islands (CK)
Record
Morgan, 2000
POLYDECTINAE
Lybia leptochelis (Zehntner,
1894)*CI
This paper
CI2-D08
Lybia tessellata (Latreille, in
Milbert, 1812)
Calman, 1909 (as Melia
tessellata); Tweedie, 1947;
Morgan, 2000; Davie, 2002
CI3-D01, CI3-D05
Wood Jones,
1909 (as Melia
tessellata);
Tweedie, 1950;
Davie, 2002
Tweedie, 1950;
Davie, 2002
Polydectus cupulifer
(Latreille, in Milbert, 1812)
XANTHINAE
Lachnopodus bidentatus (A.
Milne-Edwards, 1867)
Calman, 1909 (as Xantho
bidentatus); Ward, 1934
(as Lioxantho laevidorsalis
Miers); Davie, 2002
CI2-17, CI3-14
Tweedie, 1950
(as Paraxanthias
gibsonhilli
Tweedie); Davie,
2002
Lachnopodus gibsonhilli
(Tweedie, 1950)
298
CK1-02
RAFFLES BULLETIN OF ZOOLOGY 2014
Species Current Name
(sensu Ng et al., 2008)
Christmas Island (CI)
Record
Recent CI Stations
(2010–2012)
Lachnopodus ponapensis
(Rathbun, 1907)
Ward, 1934 (as Paraxanthias
haematostictus Ward);
Tweedie, 1947 (as
Paraxanthias ponapensis);
Morgan, 2000; Davie, 2002
CI1-13(01), CI2-09,
CI2-17, CI2-22,
CI3-23
Lachnopodus subacutus
(Stimpson, 1858)
Ward, 1934 (as Lioxantho
subacuta); Tweedie, 1947;
Morgan, 2000; Davie, 2002
CI2-17, CI2-D02,
CI3-D04
Cocos (Keeling)
Islands (CK)
Record
Recent CK Stations
(2010–2012)
Tweedie, 1950
Tweedie, 1950;
Davie, 2002
Lachnopodus tahitensis De
Man, 1889
Leptodius nudipes (Dana,
1852)
Ward, 1934; Morgan, 2000
CI1-09,
CI2-22,
CI3-14,
CI3-23,
CI1-13,
CI3-13,
CI3-16,
Ci3-25
Tweedie, 1950 (as
Xantho nudipes)
Leptodius planus Ward, 1934
Ward, 1934; Balss, 1938 (as
Xantho (Leptodius) gracilis
Dana); Tweedie, 1947 (as
Leptodius gracilis Dana);
Morgan, 2000 (as Leptodius
gracilis Dana)
CI1-09, CI3-13
Tweedie, 1950 (as
Xantho gracilis)
CK1-08, CK2-21
Leptodius sanguineus (H.
Milne-Edwards, 1834)
Calman, 1909; Balss, 1934;
Ward, 1934; Tweedie, 1947;
Morgan, 2000
CI1-09, CI1-13,
CI1-31, CI203(012), CI209(033), CI2-17,
CI2-22, CI3-07,
CI3-13, CI3-16,
CI3-23, CI3-25
Wood Jones (1909);
Tweedie, 1950 (as
Xantho sanguineus)
CK1-08, CK1-16,
CK2-08, CK2-09,
CK2-19, CK2-21,
Golf Course beach,
Trannies beach
Lioxanthodes alcocki Calman,
1909
Calman, 1909; Tweedie, 1947
(as Paraxanthias alcocki);
Morgan, 2000; Davie, 2002
CI3-16, CI3-17
Macromedaeus crassimanus
(A. Milne-Edwards, 1867)*CI
This paper
CI3-15
Macromedaeus nudipes (A.
Milne-Edwards, 1867)
Balss, 1934 (as Medaeus
nudipes); Morgan, 2000;
Davie, 2002
CI1-31(182), CI217, CI3-14, CI3-23,
CI3-25
Tweedie, 1950 (as
Medaeus nudipes);
Davie, 2002
CK1-02, CK2-17
Macromedaeus
quinquedentatus (Krauss,
1843)*CI
This paper
CI3-23
Nanocassiope alcocki
(Rathbun, 1902)
Morgan, 2000
CI3-D02, CI3-D08
Nanocassiope tridentata
Davie, 1995*CK
This paper
CK2-12
Neoxanthias impressus
(Latreille, in Milbert, 1812)
Tweedie, 1950
CK1-02
Paraxanthias aff. elegans*CI
This paper
CI3-D02
Paraxanthias notatus (Dana,
1852)
Calman, 1909 (as Xanthodes
notatus); Tweedie, 1947;
Morgan, 2000
CI1-31, CI2-17,
CI3-23
Xanthias cherbonnieri
Guinot, 1964*CI
This paper
CI2-D18
299
Mendoza et al.: Xanthidae from Christmas and Cocos Islands
Species Current Name
(sensu Ng et al., 2008)
Christmas Island (CI)
Record
Xanthias lamarckii (H. MilneEdwards, 1834)
Calman, 1909 (as Xanthodes
lamarckii); Ward, 1934;
Tweedie, 1947; Morgan,
2000)
Xanthias cf. latifrons (De
Man, 1887)
Morgan, 2000
Recent CI Stations
(2010–2012)
CI1-31,
CI2-17,
CI3-13,
CI3-16,
CI3-23,
Cocos (Keeling)
Islands (CK)
Record
Recent CK Stations
(2010–2012)
CI2-09,
CI2-22,
CI3-14,
CI3-17,
CI3-25
Wood Jones, 1909
(as Lioxantho
punctatus)
Xanthias punctatus (H. MilneEdwards, 1834)
ZOSIMINAE
Atergatis dilatatus De Haan,
1835*CI
This paper
CI3-D04
Atergatis granulatus De
Man, 1889*CI
This paper
CI2-D05
Atergatis latissimus (H.
Milne-Edwards, 1834)
Tweedie, 1947; Morgan,
2000; Davie, 2002
CI1-31
?Atergatis tweediei Ward,
1934
Ward, 1934; Tweedie, 1947;
Morgan, 2000; Davie, 2002
(as Atergatopsis tweediei)
Atergatopsis signata (Adams
& White, 1849)
Morgan, 2000
Tweedie, 1950;
Davie, 2002
Lophozozymus dodone
(Herbst, 1801)
Calman, 1909; Tweedie,
1947; Morgan, 2000; Davie,
2002
Tweedie, 1950
Tweedie, 1950;
Davie, 2002
Lophozozymus pulchellus A.
Milne-Edwards, 1867
Platypodia anaglypta (Heller,
1861)*CI
CK1-02
This paper
CI3-15
Tweedie, 1950;
Davie, 2002
Platypodia cristata (A. MilneEdwards, 1865)
Tweedie, 1950 (as
Platypodia keelingi
Tweedie); Davie,
2002
CK1-22
CI2-D18
This paper
CK2-18
Calman, 1909 (as Zozymus
aeneus); Tweedie, 1947 (as
Zozymus aeneus); Morgan,
2000; Davie, 2002
CI1-13, CI325 (typical
colouration)
Tweedie, 1950 (as
Zoozymus aeneus);
Davie, 2002
CK1-02, CK108, CK2-17
(bicoloured)
Calman, 1909 (as Leptodius
cavipes); Ward, 1934 (as
Leptodius cavipes); Tweedie,
1947 (as Leptodius cavipes);
Morgan, 2000; Davie, 2002;
Lasley & Ng, 2013
CI1-31, CI2-09,
CI2-17, CI2-22
Platypodia granulosa
(Rüppell, 1830)
Davie, 2002
Zosimus actaeoides (A.
Milne-Edwards, 1867)*CK
Morgan, 2000
Zosimus aeneus (Linnaeus,
1758)
Zozymodes cavipes (Dana,
1852)
Zozymodes pumilus (Hombron
& Jacquinot, 1846)
Tweedie, 1950
(as Zoozymodes
pumilus); Lasley &
Ng, 2013
Zozymodes xanthoides
(Krauss, 1843)*CK
This paper
300
CK1-02, CK2-09