On the identities of Sesarma obesum Dana, 1851,
and Sesarma eydouxi H. Milne Edwards, 1853
(Crustacea, Decapoda, Brachyura, Sesarmidae)
Peter K. L. NG
Department of Biological Sciences, National University of Singapore,
Kent Ridge, Singapore 119260 (Republic of Singapore)
peterng@nus.edu.sg
Christoph D. SCHUBART
Biologie 1, Universität Regensburg,
93040 Regensburg (Germany)
christoph.schubart@biologie.uni-regensburg.de
Ng P. K. L. & Schubart C. D. 2003. — On the identities of Sesarma obesum Dana, 1851,
and Sesarma eydouxi H. Milne Edwards, 1853 (Crustacea, Decapoda, Brachyura,
Sesarmidae). Zoosystema 25 (3) : 425-437.
KEY WORDS
Crustacea,
Decapoda,
Brachyura,
Sesarmidae,
Sesarma obesum,
Sesarma eydouxi.
ABSTRACT
The identities of two poorly known species of Indo-West Pacific sesarmid
crabs, which have been referred to the genus Chiromantes Gistel, 1848, are
clarified. Sesarma obesum Dana, 1851, is shown to be a senior synonym of
Metasesarma rousseauxi H. Milne Edwards, 1853. A neotype from northern
Borneo is designated to fix the identity of the species. Sesarma eydouxi H.
Milne Edwards, 1853, a species supposedly described from Vietnam and not
reported there or in the region since its description, is shown to be a junior
synonym of the Atlantic species Sesarma rectum Randall, 1840; its original
provenance almost certainly being incorrect.
MOTS CLÉS
Crustacea,
Decapoda,
Brachyura,
Sesarmidae,
Sesarma obesum,
Sesarma eydouxi.
RÉSUMÉ
Identités de Sesarma obesum Dana, 1851, et Sesarma eydouxi H. Milne
Edwards, 1853 (Crustacea, Decapoda, Brachyura, Sesarmidae).
Les identités de deux espèces peu connues de crabes Sesarmidae de l’Indoouest Pacifique, qui ont été rattachées au genre Chiromantes Gistel, 1848,
sont clarifiées. Il est démontré que Sesarma obesum Dana, 1851, est un synonyme antérieur de Metasesarma rousseauxi H. Milne Edwards, 1853. Un néotype du nord de Bornéo est désigné pour fixer l’identité de l’espèce. Sesarma
eydouxi H. Milne Edwards, 1853, espèce prétendument décrite du Viet-Nam
et non revue dans cette région depuis sa description, est démontrée être un
synonyme plus récent de Sesarma rectum Randall, 1840 de l’Atlantique ; sa
provenance originale étant presque certainement incorrecte.
ZOOSYSTEMA • 2003 • 25 (3) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris.
www.zoosystema.com
425
Ng P. K. L. & Schubart C. D.
INTRODUCTION
The identities of two Indo-West Pacific sesarmid crab species have been particularly difficult. Sesarma obesum Dana, 1851 was originally
described from Balabac off northern Borneo,
but the brevity of the description and small size
of the original figure has made its precise identity and proper generic assignment difficult to
ascertain. The type material of this species is no
longer extant.
Sesarma eydouxi H. Milne Edwards, 1853 was
described from Cochin-China (probably present day Vietnam) and was redescribed by de
Man (1892). Tesch (1917) also redescribed and
figured this species, assigning it to the genus
Holometopus H. Milne Edwards, 1853 (=
Chiromantes Gistel, 1848, fide Holthuis 1977).
Serène (1968) followed Tesch (1917) and listed
it under Holometopus (= Chiromantes). The species, however, has never been collected during
the last century and has several characters
which are atypical of Indo-West Pacific sesarmid species.
In this paper, the identities and proper systematic placements of the two species are resolved. Sesarma obesum Dana, 1851 is shown to be
a senior synonym of Metasesarma rousseauxi
H. Milne Edwards, 1853. Sesarma eydouxi H.
Milne Edwards, 1853 is a junior subjective
synonym of the Atlantic Sesarma rectum
Randall, 1840, and the original locality data
associated with the type is almost certainly
incorrect. Measurements provided (in mm)
are of the carapace width and length, respectively.
Specimens examined are deposited in the
Muséum national d’Histoire naturelle, Paris
(MNHN); Academy of Natural Sciences of
Philadelphia (ANSP); Zoological Reference
Collection of the Raffles Museum, National
University of Singapore (ZRC); Senckenberg
Museum, Frankfurt am Main (SMF); Museum
of Comparative Zoology, Harvard University,
Cambridge (MCZ); and US National Museum
of Natural History, Smithsonian Institution,
Washington DC (USNM).
426
SYSTEMATICS
Family SESARMIDAE Dana, 1851
Genus Metasesarma H. Milne Edwards, 1853
Metasesarma obesum (Dana, 1851) n. comb.
(Figs 1-3)
Sesarma obesum Dana, 1851: 252; 1852: 356, pl. 22,
fig. 10.
Metasesarma rousseauxi H. Milne Edwards, 1853: 188;
1854: 158, pl. 10, figs 1, 1a-1c. — Crosnier 1965: 73,
figs 116-120. — Ng & Davie 1995: 39.
Metasesarma granularis Heller, 1862: 522.
Metasesarma rugulosa Heller, 1865: 65.
Sesarma obesa – de Man 1887: 643.
Sesarma (Holometopus) obesa – Tesch 1917: 179.
Sesarma (Holometopus) ?obesum – Serène 1968: 107.
Chiromantes obesum – Ng & Liu 1999: 229.
M A T E R I A L E X A M I N E D . — Taiwan. Pingtung,
22.V.1998, coll. H.-C. Liu, 1 웧 , 2 웨 웨 (ZRC
1998.399); Pingtung, 19.V.1998, coll. H.-C. Liu, 1 웧
(ZRC 1999.552). — Little Liu-Kiu Island,
1.VIII.1994, coll. H.-C. Liu, C. H. Wang, 1 웧,
2 웨웨 (ZRC 1998.479). — Feng Chui-Sa, Pingtung,
24.VI.2002, coll. P. K. L. Ng, 1 웧 (ZRC).
Guam. Pago Bay, 15-18.IV.2000, coll. P. K. L. Ng, C.
H. Wang, 3 웧 웧 , 16 웨 웨 (ZRC 2000.583). —
Haputo, VIII.2001, coll. P. K. L. Ng, 2 웧웧, 1 웨
(ZRC 2001.744). — Dadi Beach, 31.VII.2001, coll.
P. K. L. Ng, 2 웧웧, 1 ovigerous 웨 (ZRC 2001.749).
Tahiti. East of Papeete, 21.XI.1996, coll. G. Paulay,
1 웨 (ZRC 2000.582); X.2001, coll. H.-C. Liu,
2 웧웧, 1 웨 (ZRC).
Australia, Christmas Island. 1939, coll. C. A. GibsonHill, 3 웧웧, 8 웨웨 (ZRC 1966.12.2.1-10).
Malaysia, Sabah. Pulau Manukan, northern Sabah,
22.VI.2000, coll. C. D. Schubart, 1 웧, 1 웨 (ZRC
2000. 1656, 1657).
Indonesia. Pangandaran, 3.III.1963, coll. R. Serène,
9 웧웧, 4 웨웨 (ZRC 1971.11.1.1-5), 1 웧 (ZRC
1971.5.3.1), 1 웧 (ZRC 1971.11.1.6). — Citerjun,
Ujung Kulon, West Java, in leaf litter, 5 m from the
shore, 30.VII.1993, coll. C. Stewart, 1 웧 15.2 ×
13.3 mm, 1 웨 15.3 × 13.4 mm (ZRC).
REMARKS
Dana (1851) described Sesarma obesum from a
specimen originating from an island in Balabac
Strait north of Borneo. Serène (1968: 107) in
listing this species under Chiromantes (as
ZOOSYSTEMA • 2003 • 25 (3)
Two species of Sesarma (Crustacea, Decapoda)
FIG. 1. — Sesarma obesum Dana, 1851, original figure, after Dana (1852: pl. 22, fig. 10a).
Holometopus) questioned its systematic position
and indicated that it might be referred to
Metasesarma. The original figure provided of
Sesarma obesum by Dana (1852: pl. 22, fig. 10) is
good (Fig. 1) and confirms that the species
should be assigned to Metasesarma. The form of
the carapace and legs strongly suggests that
Sesarma obesum Dana, 1851, is identical to
Metasesarma rousseauxi H. Milne Edwards, 1853.
This species is often reported from islands with
wide gravel or sandy beaches without rivers or
freshwater sources, as is typical for the islands
north of Borneo. Numerous specimens of this
species from the Indo-West Pacific have been
examined, and while the colour varies according
to the substrate, the lateral carapace margins of
this species are typically straight and subparallel,
giving the carapace a squarish appearance. In larger specimens (e.g., the proposed neotype),
however, the margins are gently convex.
Specimens identified as Metasesarma rousseauxi
from an island north of Sabah, relatively close to
the type locality of S. obesum, the Balabac Islands
(not easily accessible nowadays due to political
ZOOSYSTEMA • 2003 • 25 (3)
problems) were examined. The material is comparable to the specimen figured by Dana (1852).
Therefore, a male specimen from the island off
Sabah was selected as the neotype of Sesarma
obesum, thereby fixing the identity of this poorly
known species. The neotype is a male measuring
14.3 × 13.5 mm (ZRC 2000.1656), collected
from amongst supralittoral coral rubble on Pulau
Manukan, northern Sabah, East Malaysia,
Borneo, on 22 June 2000 (Figs 2; 3). This action
effectively synonymises Metasesarma rousseauxi
with Sesarma obesum, with the latter name having
priority. Furthermore, Sesarma obesum is transferred to Metasesarma.
On the basis of the identification by the late
Raoul Serène, Hartnoll (1975: 308, 322) referred
specimen(s) from Tanzania with doubt to
“Holometopus obesus”. The whereabouts of these
specimens is not known (R. G. Hartnoll pers.
comm.). Whether they are what is here identified
as Metasesarma obesum n. comb. (and M. rousseauxi) cannot be ascertained. It is interesting to
note, that while the type locality of M. rousseauxi
is the east coast of Africa (from where it has also
427
Ng P. K. L. & Schubart C. D.
A
B
C
FIG. 2. — Metasesarma obesum (Dana, 1851) n. comb., neotype, 웧 14.3 × 13.5 mm (ZRC 2000.1656), Pulau Manukan, Sabah; A,
overall view; B, carapace; C, frontal view.
428
ZOOSYSTEMA • 2003 • 25 (3)
Two species of Sesarma (Crustacea, Decapoda)
A
B
C
FIG. 3. — Metasesarma obesum (Dana, 1851) n. comb., neotype, 웧 14.3 × 13.5 mm (ZRC 2000.1656), Pulau Manukan, Sabah;
A, ventral view; B, left chela, outer view; C, left chela, inner view.
ZOOSYSTEMA • 2003 • 25 (3)
429
Ng P. K. L. & Schubart C. D.
been reported from by Crosnier [1965] and
Guinot [1967], Hartnoll (1975: 307) did not
find this species in Tanzania, but instead reported “Holometopus obesus” from the supralittoral
area of a rocky shore. This is precisely the preferred habitat of M. rousseauxi (see Ng & Davie
1995). Consequently, Hartnoll’s material is
considered to be conspecific with what is here
defined as Metasesarma obesum n. comb.
In any case, the taxonomy of what has been called
Metasesarma rousseauxi (type locality: Zanzibar),
may not be systematically stable (see Tesch 1917:
212; Crosnier 1965: 73; Ng & Davie 1995: 39,
for complete synonymy). Metasesarma granularis
Heller, 1862, and M. rugulosa Heller, 1865, are
both regarded as junior synonyms of M. rousseauxi (see Crosnier 1965; Ng & Davie 1995)
and thus now of M. obesum n. comb., but considering the extensive range of the species, it is possible that this might be a species complex with
more than one species involved. As such,
Metasesarma rousseauxi may yet prove to be different from Sesarma obesum. But on current knowledge, there is little evidence to justify separating
these two taxa. As a result, the well known supralittoral Metasesarma rousseauxi from the IndoWest Pacific should now be known as
Metasesarma obesum (Dana, 1851) n. comb. One
other Metasesarma is known from the Indo-West
Pacific, M. aubryi (A. Milne Edwards, 1869), a
species that normally occurs further inland in
forests or close to freshwater habitats (Ng &
Davie 1995).
Genus Sesarma Say, 1817
Sesarma rectum Randall, 1840
(Figs 4-7)
Sesarma recta Randall, 1840: 123.
Sesarma eydouxi H. Milne Edwards, 1853: 184. —
Heller 1865: 64. — de Man 1880: 23; 1887: 643;
1892: 248.
Sesarma müllerii A. Milne Edwards, 1869: 29.
Sesarma (Holometopus) recta – Tesch 1917: 190.
Sesarma (Holometopus) eydouxi – Tesch 1917: 150, figs
3, 4. — Serène 1968: 107.
430
Sesarma (Holometopus) rectum – Rathbun 1918: 298,
pl. 82. — Holthuis 1959: 243, fig. 61, pl. 11,
fig. 4. — Chace & Hobbs 1969: 182, fig. 62j.
Sesarma (Sesarma) rectum – von Hagen 1978: 45,
figs 1a, 2a, 3a, 4a, 5a.
Sesarma rectum – von Hagen 1975: 301, fig. 7. —
Abele 1992: 15, figs 3b, 4b, 5d, 9. — Schubart et al.
1999: 537.
Chiromantes eydouxi – Ng & Liu 1999: 229.
MATERIAL EXAMINED. — Surinam. Coll. Hering, 1 웧
32.8 × 26.7 mm, holotype of Sesarma recta Randall,
1840 (ANSP 3976). — Coquette collections,
12.V.1957, coll. L. B. Holthuis, 1 웨 (USNM
103276).
“Cochin-China”. 1 웧 31.2 × 27.2 mm, lectotype of
Sesarma eydouxi H. Milne Edwards, 1853 (MNHN
B10930).
Grenada. Fort Jeudy, 15.VIII.1994, coll. R. Diesel,
D. Horst, C. Schubart, 1 웧 21.1 × 18.4 mm, 2 웨웨
18.0 × 15.5, 16.8 × 14.5 mm (SMF 23248), 1 웨 16.5
× 14.3 mm (SMF 23249).
Brazil. Rio Grande do Norte, 1 웧 29.9 × 26.8 mm
(MCZ 6230). — Rio Paraiba do Norte, 3 웧웧 26.4 ×
22.9 mm, 17.7 × 15.1 mm, 11.7 × 9.8 mm, 1 웨
21.1 × 17.9 mm (MCZ 6231). — Pernambuco,
3 juvenile 웧웧 14.4 × 12.5 mm, 8.0 × 7.0 mm, 7.4 ×
6.4 mm, 1 웨 22.8 × 19.1 mm (MCZ 6232). — Sta
Clara, 2 웧웧 30.8 × 27.5 mm, 23.2 × 20.2 mm, 2 웨웨
29.4 × 26.1 mm, 27.8 × 24.4 mm (MCZ 6233). —
Rio de Janeiro, 4 웧 웧 26.0 × 22.0 mm, 29.7 ×
25.5 mm, 31.5 × 27.3 mm, 35.4 × 30.4 mm (MCZ
6234). — Florianopolis, 1 웧 22.6 × 19.7 mm (MCZ
6235). — Branner-Agassiz Expedition, 1.VIII.1899,
1 웧, 1 웨 (USNM 25711). — Santos, Pissaquera,
VI.1913, coll. H. Luederwaldt, 2 웧 웧 (USNM
47859). — Rio de Janeiro, Terra de Masahe, I.1912,
coll. E. Garbe, 2 웧웧 (USNM 47862). — Santos, Ilha
Casquerinita, VI.1913, coll. H. Luederwaldt, 2 juvenile
웨웨 (USNM 47867). — São Sebastião, 5 웧웧, 7 웨웨
(USNM 70986). — São Francisco, 31.X.1925, coll.
W. L. Schmitt, 4 웧웧, 1 웨 (USNM 71168). — São
Paulo, Ubatuba, mangrove at Praia Dura,
30.VIII.1992, coll. F. L. M. Mantelatto, 2 웧웧 31.2 ×
27.5 mm, 26.0 × 22.8 mm (ZRC 2000.1780). — São
Paulo, Ubatuba, mangrove at Praia Dura, III.1997,
coll. F. L. M. Mantelatto, 1 웧 22.6 × 19.9 mm (SMF
25169).
REMARKS
Sesarma eydouxi was described from CochinChina but the number of specimens actually available to H. Milne Edwards is not clear. In the
MNHN is a type specimen which is here designated as the lectotype of the species (Figs 4; 5).
ZOOSYSTEMA • 2003 • 25 (3)
Two species of Sesarma (Crustacea, Decapoda)
A
B
C
FIG. 4. — Sesarma eydouxi H. Milne Edwards, 1853, lectotype 웧 31.2 × 27.2 mm (MNHN B10930); A, dorsal view; B, carapace;
C, left chela, dorsal view.
ZOOSYSTEMA • 2003 • 25 (3)
431
Ng P. K. L. & Schubart C. D.
A
B
C
FIG. 5. — Sesarma eydouxi H. Milne Edwards, 1853, lectotype 웧 31.2 × 27.2 mm (MNHN B10930); A, frontal view; B, right chela,
dorsal view; C, left chela, dorsal view.
432
ZOOSYSTEMA • 2003 • 25 (3)
Two species of Sesarma (Crustacea, Decapoda)
A
B
C
FIG. 6. — Sesarma rectum Randall, 1840, holotype 웧 32.8 × 26.7 mm (ANSP 3976); A, overall view; B, carapace; C, front and
chelipeds, dorsal view.
ZOOSYSTEMA • 2003 • 25 (3)
433
Ng P. K. L. & Schubart C. D.
A
B
C
D
FIG. 7. — Sesarma rectum Randall, 1840, 웧 22.6 × 19.9 mm (SMF 25169); A, front, dorsal view; B, frontal view; C, right cheliped,
dorsal view; D, right chela, outer view.
434
ZOOSYSTEMA • 2003 • 25 (3)
Two species of Sesarma (Crustacea, Decapoda)
Heller (1865) subsequently reported this species
from Madras (India), and de Man (1880) recorded it on the basis of specimens from an unknown
provenance (measurements of largest male 33.0 ×
27.0 mm), and he subsequently redescribed the
type (de Man 1892). Tesch (1917) redescribed in
detail the specimens of de Man (1880), and made
a telling comment: “This species is exceedingly
alike Ses. recta Randall of America, and I own to
be at a loss to indicate any important point of difference between the two species” (Tesch 1917:
153).
Sesarma rectum was described from Surinam, and
S. muellerii (type locality: Brazil, 30.0 by 27.0
mm) is now generally regarded as its junior synonym (for detailed synonymy, see Tesch 1917:
190; Abele 1992: 15; for recent distribution
records, see Schubart et al. 1999). Photographs of
Sesarma rectum, a dried type specimen in the
Philadelphia Museum (Fig. 6), and S. eydouxi
(Figs 4; 5) were compared, and only minimal differences were observed, confirming that the two
species are synonymous. The holotype of Sesarma
rectum has chelae with 15 more or less evenly
spaced dactylar tubercles, whereas the lectotype
of S. eydouxi has 14 (Figs 4C; 5B, C; 6B, C). In
both cases, the tubercles have corneous tips
which are slightly pointing distally (see also Abele
1992: fig. 4b with only 12 tubercles; fig. 9 with
14 tubercles, 14-16 tubercles in text). The upper
surface of the chelar palm of both type specimens
has a continuous ridge of unpectinated tubercles
(Figs 4C; 5C; 6C; 7C), a typical character for the
American species of Sesarma s.s. (see von Hagen
1978; Abele 1992: fig. 2e). The chelar carpus of
both types is characterised by a triangular tooth
at the inner distal corner and a coarsely granulated outer surface (Figs 4C; 7C). The lectotype of
S. eydouxi has a slight indication of an anterolateral tooth, without forming a real notch or
tooth (Fig. 4A, B). In the original description, H.
Milne Edwards (1853: 184) refers to it as:
“Carapace rugueuse, pubescente et obscurément
bidentée de chaque côté”. The indicated tooth is
not as evident in the type of S. rectum (Fig. 6B),
but also known from this species: “Slight emargination present, indicated by ridge below angle
ZOOSYSTEMA • 2003 • 25 (3)
with second stronger ridge below first” (Abele
1992: 15, fig. 9). The types of S. rectum and S.
eydouxi also agree in the presence of two rows of
setae delimiting the epistomial Verwey’s groove
(Figs 5A; 7B) (see von Hagen 1978; Abele 1992:
fig. 2d), in the shape of the frontal lobes, and in
the particularly broad meri of the walking legs
(Figs 4A; 6A) (meri of fourth walking leg less
than twice as long as wide). Based on the measurements taken for us in Paris and Philadelphia,
the carapace shapes of the two type specimens
appeared to differ substantially. Maximum carapace width divided by maximum carapace length
resulted in a ratio of 1.23 for S. rectum and 1.15
for S. eydouxi. Re-measurement of the photographed carapaces, however, revealed a ratio of
1.13 for both types. The measurements obtained
by de Man (1892) for the type of S. eydouxi
(31.25 × 26.25 mm) resulted in a ratio of 1.19.
The discrepancies in the original ratio for S. rectum could be because the width and/or length
measurements were taken in a different manner
by the ANSP staff.
In summary, the two species are here regarded as
synonymous and that the original locality data of
S. eydouxi must be incorrect. In fact, numerous
collections from around that area and Southeast
Asia in general have never uncovered a species
even resembling S. rectum. The record of
“S. eydouxi” from Madras (India) by Heller (1865:
64) will need to be rechecked – it is not S. eydouxi
as presently defined. As indicated by von Hagen
(1978) and genetically confirmed by Schubart et
al. (1998a), S. rectum clearly belongs to Sesarma
s.s., despite the absence of an anterolateral tooth.
The genus Sesarma presently contains 16 species,
all from the American Atlantic and eastern Pacific
(Abele 1992; Schubart et al. 1998b).
Acknowledgements
The authors are indebted to Tan Swee Hee for
checking and photographing the specimens in
the MNHN and to Danièle Guinot for her kind
help. Lu Eldredge helped with the original plates
from Dana (1852), which were important for
identifying the identity of Sesarma obesum. Ardis
435
Ng P. K. L. & Schubart C. D.
Johnston (MCZ) and Karen Reed (USNM)
loaned material of Sesarma rectum from their respective museums. Elana Benamy, Gary Rosenberg and Albert M. Greenfield (Digital Imaging
Center of the Academy of Natural Sciences of
Philadelphia) are being thanked for making available measurements and digital photographs of
the holotype of Sesarma rectum.
REFERENCES
ABELE L. G. 1992. — A review of the grapsid crab
genus Sesarma (Crustacea: Decapoda: Grapsidae) in
America, with the description of a new genus.
Smithsonian Contributions to Zoology 527: 1-60.
CHACE F. A. JR. & HOBBS H. H. JR. 1969. — The
freshwater and terrestrial decapod crustaceans of the
West Indies with special reference to Dominica.
Bulletin of the United States National Museum 292:
1-258.
C R O S N I E R A. 1965. — Crustacés Décapodes,
Grapsidae et Ocypodidae. Faune de Madagascar 18:
1-143, pls 1-11, text-figs 1-260.
DANA J. D. 1851. — Conspectus crustaceorum quae in
orbis terrarum circumnavigatione, Carolo Wilkes e
classe Reipublicae Foederatae Duce, lexit et descripsit
J. D. Dana. Crustacea Grapsoidea, (Cyclometopa,
Edwardsii). Proceedings of the Academy of Natural
Sciences of Philadelphia 5: 247-254.
DANA J. D. 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. 13 (2): 686-1618. C. Sherman,
Philadelphia. Atlas [1855]. Vol. 14, 27 p., pls 1-96.
G ISTEL J. N. F. X. 1848. — Naturgeschichte des
Tierreichs für höhere Schulen. Stuttgart, xvi + 216 p.,
pls 1-32.
G U I N O T D. 1967. — La faune carcinologique
(Crustacea Brachyura) de l’océan Indien occidental
et de la mer Rouge. Catalogue, remarques biogéographiques et bibliographie, in Réunion de
Spécialistes C.S.A. sur les Crustacés, Zanzibar 1964.
Mémoires de l’Institut fondamental d’Afrique Noire,
Ifan-Dakar (77), 1966 (1967): 237-352.
HAGEN H.-O. VON 1975. — Klassifikation und phylogenetische Einordnung der Lautäußerungen von
Ocypodiden und Grapsiden (Crustacea,
Brachyura). Zeitschrift für Systematik und
Evolutionsforschung 13: 300-316.
HAGEN H.-O. VON 1978. — The systematic position
of Sesarma (Sesarma) rectum Randall and a new definition of the subgenus Sesarma (Brachyura,
Grapsidae). Crustaceana 34 (1): 45-54.
436
H A R T N O L L R. G. 1975. — The Grapsidae and
Ocypodidae (Decapoda: Brachyura) of Tanzania.
Journal of Zoology 177: 305-328.
H ELLER C. 1862. — Neue Crustaceen gesammelt
während der Weltumseglung der K. K. Fregatte
Novara. Verhandlungen der kaiserlich-königlichen zoologisch-botanischen Gesellschaft in Wien 12: 519-528.
H ELLER C. 1865. — Crustaceen, in Reise der
Oesterreichischen Fregatte Novara um die Erde in
den Jahren 1857, 1858, 1859 unter den Befehlen
des Commodores B. von Wüllerstorf-Urbair.
Zoologischer Theil. Zweiter Band 3 (1): 1-280, 25 pls.
HOLTHUIS L. B. 1959. — The Crustacea Decapoda of
Suriname (Dutch Guiana). Zoologische Verhandelingen 44: 1-296, pls 1-11.
HOLTHUIS L. B. 1977. — The Grapsidae, Gecarcinidae
and Palicidae (Crustacea: Decapoda: Brachyura) of
the Red Sea. Israel Journal of Zoology 26: 141-192.
MAN J. G. DE 1880. — On some species of the genus
Sesarma Say and Cardisoma Latr. Notes Leyden
Museum 2 (5): 21-36.
MAN J. G. DE 1887. — Übersicht der Indo-pacifischen Arten der Gattung Sesarma Say nebst einer
Kritik der von W. Hess und E. Nauck in den Jahren
1865 und 1880 beschriebenen Decapoden.
Zoologische Jahrbücher, Abtheilung für Systematik,
Geographie und Biologie der Thiere 2: 639-722.
MAN J. G. DE 1892. — Carcinological studies in the
Leyden Museum, No. 6. Notes Leyden Museum 14
(36): 225-264, pls 7-10.
MILNE EDWARDS A. 1869. — Notes sur quelques nouvelles espèces du genre Sesarma (Say). Nouvelle
Archives du Muséum d’Histoire naturelle 5: 25-31.
MILNE EDWARDS H. 1853. — Mémoires sur la famille
des Ocypodiens, suite. Annales de Sciences naturelle,
Zoologie (Paris) sér. 3 (Zoologie) 20: 163-228, pls
6-11 [reprinted in the undated Mélanges carcinologiques: 129-196].
M ILNE E DWARDS H. 1854. — Notes sur quelques
Crustacés nouveaux ou peu connus conservés dans la
collection du Muséum d’Histoire naturelle. Archives
du Muséum d’Histoire naturelle 7: 145-192, pls 9-16
[reprinted in the undated Mélanges carcinologiques].
NG P. K. L. & DAVIE P. J. F. 1995. — The terrestrial
sesarmine crabs of the genera Metasesarma and
Geosesarma (Crustacea: Decapoda: Brachyura:
Grapsidae) of Ujung Kulon, West Jawa, Indonesia.
Tropical Biodiversity 3: 29-43.
NG P. K. L. & LIU H.-C. 1999. — The taxonomy of
Sesarma tangi Rathbun, 1931 and S. stormi de Man,
1895 (Crustacea: Decapoda: Brachyura: Grapsidae:
Sesarminae), with establishment of a new genus for
S. stormi. Zoological Studies 38 (2): 228-237.
RANDALL J. W. 1840. — Catalogue of the Crustacea
brought by Thomas Nuttall and J. K. Townsend,
from the west coast of North America and the
Sandwich Islands, with descriptions of such species as
are apparently new, among which are included several species of different localities, previously existing in
ZOOSYSTEMA • 2003 • 25 (3)
Two species of Sesarma (Crustacea, Decapoda)
the collection of the Academy. Journal of the Academy
of Natural Science of Philadelphia 8: 106-147, pls 1-5.
R ATHBUN M. J. 1918. — The grapsoid crabs of
America. United States National Museum Bulletin
97: 1-461, 161 pls.
SCHUBART C. D., DIESEL R. & HEDGES S. B. 1998a.
— Rapid evolution to terrestrial life in Jamaican
crabs. Nature 393: 363-365.
SCHUBART C. D., REIMER J. & DIESEL R. 1998b. —
Morphological and molecular evidence for a new
endemic freshwater crab, Sesarma ayatum sp. n.,
(Grapsidae, Sesarminae) from eastern Jamaica.
Zoologica Scripta 27 (4): 373-380.
SCHUBART C. D., HORST D. & DIESEL R. 1999. —
First record of Sesarma rectum Randall (Brachyura,
Grapsidae, Sesarminae) from the Lesser Antilles.
Crustaceana 72 (5): 537-538.
SERÈNE R. 1968. — The Brachyura of the Indo Pacific
Region, in Prodromus for a check list of the
non-planctonic marine fauna of South East Asia.
Singapore National Academy of Science Special
Publication 1: 33-112.
TESCH J. J. 1917. — Synopsis of the genera Sesarma,
Metasesarma, Sarmatium and Clistocoeloma, with a
key to the determination of the Indo-Pacific species.
Zoologische Mededelingen 3: 127-260, pls 15-17.
Submitted on 25 February 2002;
accepted on 13 August 2002.
ZOOSYSTEMA • 2003 • 25 (3)
437