THE RAFFLES BULLETIN OF ZOOLOGY 2007
THE RAFFLES BULLETIN OF ZOOLOGY 2007 Supplement No. 16: 191–226
Date of Publication: 31 Dec.2007© National University of Singapore

THE PINNOTHERID TYPE MATERIAL OF SEMPER (1880), NAUCK (1880)
AND BÜRGER (1895) (CRUSTACEA: DECAPODA: BRACHYURA)
Shane T. Ahyong
Marine Biodiversity and Biosecurity, National Institute of Water and Atmospheric Research,
Private Bag 14901, Kilbirnie, Wellington, New Zealand
Email: s.ahyong@niwa.co.nz

Peter K. L. Ng
Department of Biological Sciences, National University of Singapore,
Kent Ridge, Singapore 119260, Republic of Singapore
Email: peterng@nus.edu.sg

ABSTRACT. – The major study of pinnotherid crabs by Otto Bürger (1895) is a landmark publication forstudents of the Pinnotheridae, not only for the many new species described, but also for its introduction of
useful species characters. Bürger (1895) described and illustrated 28 new species, mainly from the Philippines,
and reported additional species including those described by his colleagues, Semper and Nauck. Unfortunately,
few subsequent workers have examined the ‘Bürger’ material–a significant problem because his species
accounts are inadequate by modern taxonomic standards leading to questionable identifications. Therefore,
all available ‘Bürger’ material has been re-examined herein. Of the 31 new species described by Bürger,
Semper and Nauck, types of 21 species are still extant. Species for which original types are still extant are
redescribed and refigured, with lectotypes usually selected. Species for which type material is lost are briefly
diagnosed and illustrated by their original figures. Species for which neotypes have been assigned by previous
studies are illustrated and diagnosed for completeness. Generic assignments of all species are reassessed andupdated. Bürger’s specimens of Pinnotheres parvulus Stimpson, 1858, are syntypes of P. alcocki Rathbun,
1909, and transferred to Arcotheres. Pinnotheres glaber Bürger, 1895, is synonymised with P. impressus
Bürger, 1895, and transferred to Orthotheres. Pinnotheres rhombifer Bürger, 1895, is synonymised with P.
latissimus Bürger, 1895, and transferred to Arcotheres. Pinnotheres rotundatus Bürger, 1895, is alsosynonymised with P. consors Bürger, 1895, and transferred to Arcotheres.The type series of Arcotheres
palaensis Bürger, 1895, comprised two species, one of which is referable to A. guinotae Campos, 2001; alectotype is selected to retain validity of both species. The type series of Xenophthalmus latifrons Bürger,
1895 (here transferred to Arcotheres), contained two species, one of which is named A. rayi, new species.
Pinnotheres gracilis Bürger, 1895, and P. burgeri Rathbun, 1909, are transferred to Viridotheres.
Miscellaneous, unstudied material in the ‘Bürger’ collection included an undescribed Viridotheres, herein
named V. otto, new species. Pinnotheres placunae Hornell & Southwell, 1909, is also referred to Arcotheres.
KEY WORDS. – Crustacea, Decapoda, Brachyura, Pinnotheridae, type material, new species, Arcotheresrayi, Viridotheres otto, taxonomy.

INTRODUTION
The paper by Otto Bürger (1895) on Indo-West PacificPinnotheridae is a landmark publication in brachyuran studies.
Bürger (1895) described and illustrated 28 new species,
mainly from the Philippines, but more importantly, introduceda suite of useful taxonomic characters. As Manning (1993a:
126) commented “Almost a century ago, Bürger (1895)
introduced a variety of characters that could be used to
distinguish species and groups of Pinnotheres, e.g., the shapeand relative lengths of the distal segments of the MXP3, theshape of the carapace, the relative dactylar and overall lengthsof the walking legs. Few workers have adopted the charactersused by Bürger, yet his study laid the groundwork for arevision of the genus by providing a suite of very distinctive
characters usable at the generic level.” Bürger (1895) had
been extensively cited by almost all pinnotherid workers but
few have examined or attempted to revise his material. Thishas been a serious problem because the descriptions andfigures provided by Bürger are inadequate by modern
taxonomic standards.
In their major synopsis of the Pinnotheridae, Schmitt et al.
(1973) located many of Bürger’s specimens as well as those191

Ahyong & Ng: Pinnotherid types of Bürger (1895)
studied by two of his associates at Germany’s University of
Göttingen, Semper and Nauck, who between them describedseveral new species. Schmitt et al. (1973) commented thatthey could not find the types of several species, which may
be lost.
In 1999, when the second author and the late Raymond
Manning embarked on a study of pinnotherids associated withsea cucumbers (see Ng & Manning, 2003), attempts weremade to trace Bürger’s, Semper’s and Nauck’s material. It
was discovered that the Smithsonian Institution held six lotsof six species obtained by C. Semper, including material used
for three important pinnotherid papers (Semper, 1880; Nauck,
1880; Bürger, 1895). Oddly, these species, Pinnotheres flavus
Nauck, 1880, P. holothuriae Semper, 1880, P. affinis Bürger,
1895, P. exiguus Bürger, 1895, P. glaber Bürger, 1895, andXenophthalmus latifrons Bürger, 1895, were not cited by
Schmitt et al. (1973), although Mary J. Rathbun was awareof the type status of these specimens based on data from theirlabels. Michael Türkay and Andreas Allspach confirmed that
the Senckenberg Museum in Frankfurt, Germany, was theeventual repository of the extant crustacean material fromthe Göttingen University, and still possessed types of two
holothurian-dwelling pinnotherid species they were studying:
Pinnotheres flavus Nauck, 1880, and Holothuriophilus
trapeziiformis Nauck, 1880 (see Ng & Manning (2003) fora detailed account). It is important here to note that although
Ng & Manning (2003) indicated that some of Bürger’s (1895)
types were present in the Zoological Museum in Berlin, we
have since been informed by Oliver Coleman (pers. comm.)
that there are actually none in the Berlin collections.
In 2001, Michael Türkay generously offered the second author
the opportunity to study the “Göttingen” collection of
pinnotherids but commented that there were many problems.
Andreas Allspach prepared a list of the Göttingen material
in the Senckenberg Museum, warning that sometime in thepast (possibly several times), some of the bottles containing
the “Göttingen” pinnotherids had dried out and were refilledmuch later. Consequently, the condition of some specimens
was poor. Moreover, some bottles had been broken, thecontents may have mixed, and not all specimens listed by
Semper (1880), Nauck (1880) and Bürger (1895) could be
accounted for. There was also material from Göttingen that
had apparently never been published on by these authors.
Nevertheless, the importance of this material for a clearer
understanding of pinnotherid taxonomy, and the fact that there
were still many extant types, compelled the complete studyof the material. In this paper, the pinnotherid species describedby Semper (1880), Nauck (1880) and Bürger (1895) are listed,
and the status of the type material summarised.
Of the 31 species described by Semper, Nauck and Bürger,
types of 21 species are still extant (see also Ng & Manning,2003). Of the 10 species for which types are lost, Ng &Manning (2003) selected neotypes for two species.

192

For species whose types are still extant, lectotypes are in mostcases selected and the taxon is redescribed and refigured.
Where type material is lost, the original figures have been
reproduced and in a brief diagnosis, the key characters are
identified for the benefit of future workers. Genericassignments are also reassessed and taxonomy updated. Of
these, five species [Pinnotheres flavus Nauck, 1880, P.
semperi Bürger, 1895 (both now in Holotheres Ng &Manning, 2003), P. holothuriae Semper, 1880, P. ortmanni
Bürger, 1895, P. tenuipes Bürger, 1895 (both now in
Buergeres Ng & Manning, 2003)] were dealt with by Ng &Manning (2003) and one species (Durckheimia caeca Bürger,1895) was redescribed by Ahyong & Ng (2005).

MATERIALS AND METHODS
Terminology used herein follows Manning (1993a). The
following abbreviations are used: MXP3 = third maxilliped;
the walking legs, pereiopods 2–5, are referred to as P2–5,
respectively; Gl = male first pleopod. Measurements are of
the carapace width and length (cw. and cl., respectively),respectively in millimetres (mm). Specimens examined aredeposited in the Senckenberg Museum, Frankfurt-am-Main(SMF); National Museum of Natural History, Smithsonian
Institution, Washington D.C. (USNM); and Zoological
Reference Collection of the Raffles Museum, National
University of Singapore (ZRC). The Göttingen material in
the SMF is catalogued separately from the main crustaceancollections. The specimens carry original Göttingen Museum
numbers (here indicated with the prefix Go) but are also
separately catalogued in SMF under the abbreviation ZMG(for Zoologisch Museum Göttingen). Both sets of registration
numbers are included in this study.
Since many of Bürger’s species have not been adequately
characterised until now, most published distribution records
will require verification. Consequently, synonymies are
restricted to the primary citation, and studies dealing directlywith Bürger’s material and/or types. A description andillustration of the type material is given for species not treatedby Ng & Manning (2003) and Ahyong & Ng (2005). For
completeness, species dealt with by Ng & Manning (2003)
and Ahyong & Ng (2005) are illustrated and briefly
diagnosed. Numerous currently used species characters suchas walking leg asymmetry, segment proportions and dactylar
setation are difficult or impossible to interpret from Bürger’s
account and figures, so it is not always possible to provide
adequate diagnoses for species lacking type material. For
species lacking types, the original figures and a brief list ofdiagnostic characters extracted from Bürger (1895) is given
as a starting point for future studies. Table 2 lists species
reported by Bürger (1895) in their original and current
combinations, along with unpublished specimens from the‘Bürger collection’, the number of specimens originally cited
and material still extant.

THE RAFFLES BULLETIN OF ZOOLOGY 2007
TAXONOMYPINNOTHERIDAE DE HAAN, 1833
Arcotheres alcocki (Rathbun, 1909), new combination(Fig. 1)
Pinnotheres parvulus – Bürger, 1895: 375–376, Pl. 9: Fig. 18, Pl.
10: Fig. 17 [not P. parvulus Stimpson, 1858].
Pinnotheres alcocki Rathbun, 1909: 114 [type locality: Burias, thePhilippines, by present lectotype designation].
Type material. – Lectotype: SMF-ZMG180 (Go655b), female (11.0
× 8.0 mm), Burias, the Philippines.

Description. – Female: Carapace subhexagonal, wider thanlong, anterior margin blunt; front slightly projecting, convex;
posterior margin straight. Eyes small, partially visible indorsal view. MXP3 with propodus about 3 times as long as
high, apex blunt, rounded; dactylus not reaching propodal
apex; ischiomerus about twice as long as wide, outer margin
convex, inner margin with proximal 3/4 slightly concave,angular at distal quarter; exopod outer margin concave, inner
margin relatively straight, flagellum two-segmented. Chelipedwith dactylus longer than half palm length; palm about twice
as long as high, mesioventral margin setose; dactylus occlusal
margin with large proximal tooth; pollex occlusal margin withsmall proximal tooth, sparsely setose. Walking legs dorsallyand ventrally unarmed; relative lengths of meri P4 > P3 > P2
> P5. P2–P5 dactyli relative lengths P5 > P4 > P3 > P2. P2
and P3 dactyli less than half propodal length. P4 asymmetricalin length; sparsely setose; left and right dactyli subequal inlength, shorter than propodus, setose ventrally. P5 dactylus

as long as propodus, slightly longer than P4 dactylus; ventrally
setose, dorsally setose distally; ventral distal margin with 2
rows of 10–12 minute spinules. Abdomen extending to buccal
region, covering bases of walking legs. Male: not known.
Remarks. – One of Bürger’s (1895) three specimens that wasreported as P. parvulus from Burias is still extant. Rathbun(1909) regarded records of P. parvulus by Bürger (1895) fromthe Philippines, de Man (1887: 105, 1888: 383) from the
Mergui Archipelago and Indonesia, and Alcock (1900) froman unspecified locality, as misidentifications, and proposeda new name for the species, P. alcocki. The relative lengthsof the walking leg dactyli in P. alcocki, in which the P4 andP5 dactyli are longer than those of P2–3 indicate thatRathbun’s species is referable to Arcotheres. Rathbun (1909)
did not designate type specimens for A. alcocki, so all of thematerial reported as P. parvulus by Bürger (1895), de Man(1887, 1888) and Alcock (1900) are syntypes. Significantly,
the Mergui Archipelago syntype of de Man (1887) andreported by Gordon (1936), although said to be in poor
condition, differs from the Philippine syntype, most notably
in the setation and relative lengths of the P4 and P5 dactyli,
with the P4 dactylus about 0.6 times rather than subequal tothat of P5. Thus, the syntypes of A. alcocki are composite.
To fix the identity of A. alcocki, Bürger’s Philippine syntype(ZMG180) is herein designated as the lectotype. Fixation of
the Philippine syntype as lectotype of A. alcocki raises thequestion of the identity of the Mergui Archipelago material.
The Mergui Archipelago material almost certainly represents
an undescribed species that is similar to, but probably distinct
from Japanese material reported as P. alcocki by Takeda &Konishi (1988), also an undescribed species. In particular,

Fig. 1. Arcotheres alcocki (Rathbun, 1909), new combination, SMF-ZMG180, lectotype female (11.0 × 8.0 mm), Burias, the Philippines:
A, dorsal habitus. B, right cheliped. C, right MXP3. Scale bars: A, B = 2.0 mm; C = 0.7 mm.

193

Ahyong & Ng: Pinnotherid types of Bürger (1895)
the propodus of MXP3 in the Japanese material tapers lessstrongly than that of the Mergui Archipelago specimen (ifthe figures are correct). Further action on the Mergui andJapanese species is deferred until specimens can be examined.
The present status of material reported by Alcock (1900) is
not known.
Takeda & Konishi (1988) compared and contrasted their
Japanese “P. alcocki” with similar congeners including P.
sinensis Shen, 1932. Ironically Pinnotheres sinensis, referable
to Arcotheres, agrees well in almost all respects with A.
alcocki sensu stricto, particularly in MXP3 morphology,
walking leg segment proportions, and the ornamentation of
the P5 dactyli including two ventral rows of minute spinules(based on Shen, 1932; Takeda & Konishi, 1988; Dai & Yang,

1991). Judging from the type description and figures, A.
sinensis differs from A. alcocki in its slightly shorter P4 dactyli(0.8 vs. 0.9 P5 length) and possibly in carapace width: length
ratio as well, 1.4 versus 1.2, based on Shen’s (1932) figure78b. Arcotheres alcocki and A. sinensis are possiblysynonymous, but assessment of the status of the latter must
await a redescription of type material.

Arcotheres arcophilus (Bürger, 1895)(Fig. 2)
Pinnotheres arcophilus Bürger, 1895: 371, Pl. 9: Fig. 10, Pl. 10:
Fig. 10 [type locality: Ubay, the Philippines, from Arca, bypresent lectotype designation].

Fig. 2. Arcotheres arcophilus (Bürger, 1895), new combination, Ubay, the Philippines: A–C, SMF-ZMG172a, female lectotype (5.3 × 4.7mm); D, E, SMF-ZMG172b, male paralectotype, (2.8 × 2.5 mm). A, D, dorsal habitus; B, right cheliped; C, right MXP3; E, abdomen. Scalebars: A = 2 mm; B, D = 1.4 mm; C, E = 0.5 mm.

194

THE RAFFLES BULLETIN OF ZOOLOGY 2007
Type material. – Lectotype: SMF-ZMG172a (Go954a), female (5.3
× 4.7 mm), Ubay, the Philippines, from Arca, C. Semper, 1863–
1864. Paralectotypes: SMF-ZMG172b (Go954a), 1 male (2.8 × 2.5mm), 1 shrivelled female (5.1 × 4.5 mm), type locality.

Description. – Female: Carapace subhexagonal, slightly wider
than long, anterior margin sharp, front faintly projecting,slightly convex. Eyes small, partially visible in dorsal view.
MXP3 with propodus about 3 times as long as high, apex
rounded; dactylus not reaching propodal apex; ischiomerus
about twice as long as wide, outer margin convex, inner
margin with proximal 3/4 slightly concave, angular at distal
quarter; exopod inner and outer margin convex, flagellum
two-segmented. Cheliped with dactylus longer than half palm
length; palm about 1.5 times as long as high, mesioventral
margin setose; dactylus occlusal margin with large proximal
tooth and short setae; pollex occlusal margin with 2 blunt
proximal teeth, sparsely setose. Walking legs dorsally andventrally unarmed; relative lengths of meri P2 = P3 > P4 >P5. P2–P5 dactyli relative lengths P5 > P4 > P3 > P2. P2 andP3 dactyli about half propodal length. P4 asymmetrical,
sparsely setose; longer dactylus longer than propodus, about
twice length of shorter dactylus. P5 dactylus distinctly longer
than propodus; ventrally setose, dorsally setose distally.
Abdomen extending to buccal region, covering bases ofwalking legs. Male: Carapace subcircular, slightly wider thanlong, front projecting, margin convex. Eyes visible in dorsal
view. MXP3 as in female. Cheliped as in female. Walking
legs dorsally and ventrally unarmed; relative lengths of meri
P2 = P3 > P4 > P5. P2–P5 dactyli relative lengths P5 > P4
> P3 = P2. P2 and P3 dactyli about 2/3 propodal length. P4
asymmetrical; longer dactylus about 0.9 propodus length,
about 1.5 times length of shorter dactylus. P5 dactylus slightly
longer than propodus. Abdomen widest at somites 2 and 3,
tapering distally to evenly rounded telson; telson wider thanlong.
Remarks. – Campos & Manning (2001) transferred P.
arcophilus to Arcotheres. All three original syntypes of A.
arcophilus are extant, of which the larger female syntype ishere designated the lectotype.

Arcotheres coarctatus (Bürger, 1895), new combination(Fig. 3A)
Pinnotheres coarctatus Bürger, 1895: 369, Pl. 9: Fig. 7, Pl. 10: Fig.
7 [type locality: Zamboanga, the Philippines, from “Cahebe”(= Polymesoda spp.)].
Type material. – Lost.

Diagnosis. – Female: Carapace subquadrate, with shallow
dorsal longitudinal groove extending from above each orbit
to about midlength. Longer P4 dactylus shorter than half
propodal length and about half-length of P5 dactylus. P5
dactylus shorter than propodus. MXP3 propodus with tapering
apex; dactylus overreaching propodal apex; ischiomerus withobtuse but distinct inner distal angle. Male: not known. [after
Bürger, 1895: 369, Pl. 9: Fig. 7, Pl. 10: Fig. 7]

Remarks. – In Pinnotheres coarctatus, the dactyli of P4–5
are longer than those of P2–3, and P4 is asymmetrical inlength from left to right. Therefore, P. coarctatus is hereintransferred to Arcotheres.
The holotype of A. coarctatus from Zamboanga is lost. Three
specimens from Singapore, however, are available (ZRC),
obtained from within the mangrove clam Polymesoda, andcorrespond well to Bürger’s (1895) account and figures of A.
coarctatus. At present, since A. coarctatus appears to bereadily recognizable and cannot be confused with any otherspecies, a neotype for the species is not necessary.

Arcotheres exiguus (Bürger, 1895)(Fig. 4)
Pinnotheres exiguus Bürger, 1895: 377, Pl. 9: Fig. 19, Pl. 10: Fig.
30 [type locality: Samar Island, the Philippines, by present
lectotype designation].
Type material. – Lectotype: USNM 32432, female (5.6 × 4.6 mm),
Samar Island, the Philippines, C. Semper, 1876.

Description. – Female: Carapace subhexagonal, wider thanlong, anterior margin well defined; front slightly projecting,transverse; posterior margin broadly convex. Eyes small,partially visible in dorsal view. MXP3 with propodus about
3 times as long as high, apex blunt, rounded; dactylus notreaching propodal apex; ischiomerus about twice as long as
wide, outer margin convex, inner margin with proximal 3/4
concave, angular at distal quarter; exopod inner and outermargin convex, flagellum one-segmented. Cheliped withdactylus about 2/3 palm length; palm about twice as long as
high, mesioventral margin setose; dactylus occlusal margin
with large proximal tooth; pollex occlusal margin with 2 small
proximal teeth, sparsely setose. Walking legs dorsally andventrally unarmed; relative lengths of meri P4 > P2 = P3 >P5. P2–P5 dactyli relative lengths P5 > P4 > P3 > P2. P2 andP3 dactyli less about half propodal length. P4 sparsely setose;left dactylus shorter than propodus; right dactylus not known.
P5 dactylus slightly long as propodus; ventrally setose,
dorsally setose distally. Abdomen extending to buccal region,
covering bases of walking legs. Male: not known.
Remarks. – Campos & Manning (2001) transferred P. exiguus
to Arcotheres. Of the four original syntypes, only one isextant, and is herein designated the lectotype. Unfortunately,
the distal three segments of the right P4 are missing, structuresthat bear diagnostic value in Arcotheres. Judging by the
relative length of the dactylus of the left P4, and the figure
given in the type account, the missing dactylus is probably
the longer.

Arcotheres latifrons (Bürger, 1895), new combination(Fig. 5)
Xenophthalmus latifrons Bürger, 1895: 387, Pl. 9: Fig. 32, Pl. 10:
Fig. 32 [type locality: Bohol, the Philippines].

195

Ahyong & Ng: Pinnotherid types of Bürger (1895)

Fig. 3. Dorsal habitus and MXP3: A, Arcotheres coarctatus (Bürger, 1895), new combination, female holotype; B, Buergeres tenuipes(Bürger, 1895), female holotype, Ubay, the Philippines; C, Nepinnotheres pectinicola (Bürger, 1895), new combination, female holotype;
D, Nepinnotheres rathbunae (Schmitt, McCain & Davidson, 1973), new combination, female holotype; E, Orthotheres laevis (Bürger, 1895),
female holotype; F, Orthotheres longipes (Bürger, 1895), new combination, female holotype. (After Bürger, 1895, pl. 9: figs 1, 7, 8, 11,
22, 25, 31, pl. 10: figs 1, 7, 8, 11, 22, 24, 31).

196

THE RAFFLES BULLETIN OF ZOOLOGY 2007
Type material. – Lectotype: USNM 32436, female (9.5 × 8.1 mm),
Mariveles or Bohol, the Philippines, coll. Semper. Paralectotype:
SMF-ZMG952, female (11.3 × 8.7 mm), Mariveles or Bohol,
Semper (not Xenophthalmus latifrons Bürger, 1895; holotype of A.
rayi, new species, see later).

Description. – Female: Carapace subtrapezoid, wider thanlong, anterior margin well defined; front not projecting but
demarcated by emargination lateral to frontal margin. Eyes
absent. MXP3 with propodus greater than 3 times as long as
high, apex blunt; dactylus not reaching propodal apex;
ischiomerus length less than twice as long as wide, outermargin convex, inner margin with proximal 3/4 slightly
concave, angular at distal quarter; exopod inner and outermargin convex, flagellum two-segmented. Cheliped withdactylus about half palm length; palm about 2.5 times as longas high, mesioventral margin setose; dactylus occlusal margin
with large proximal tooth; pollex occlusal margin with 2 small
proximal teeth, sparsely setose. Walking legs dorsally andventrally unarmed; relative lengths of meri P4 > P3 >P2 >P5. P2–P5 dactyli relative lengths P4 > P5 > P3 > P2. P2 andP3 dactyli about half propodal length. P4 asymmetrical inlength; merus half carapace length; longer dactylus shorter
than propodus, about twice length of shorter dactylus. P5
dactylus as long as propodus; ventrally setose. Abdomen
extending to buccal region, covering bases of walking legs.
Male: not known.
Remarks. – Tesch (1918) argued that the present species is
not correctly placed in Xenophthalmus. The dactyli of P4–5
are longer than those of P2–3, so X. latifrons is transferredFig. 4. Arcotheres exiguus (Bürger, 1895), USNM 32432, female (5.6 × 4.6 mm), Samar Island, the Philippines; A, dorsal habitus; B, rightcheliped; C, right MXP3. Scale bars: A = 2.0 mm; B = 1.5 mm; C = 1.0 mm.

197

Ahyong & Ng: Pinnotherid types of Bürger (1895)
to Arcotheres. Two of the original six syntypes of A. latifrons
are extant, both from Bohol, but each represent a separate
species. One syntype (USNM 32436) represents the blind
species figured by Bürger (1895) and is herein designated as
the lectotype. Thus, A. latifrons is unique in the genus forlacking eyes. The paralectotype, which represents a separate
species, is described below as new (see Arcotheres rayi, newspecies).

Arcotheres latus (Bürger, 1895), new combination(Fig. 6)
Pinnotheres latus Bürger, 1895: 374–375, Pl. 9: Fig. 16, Pl. 10:
Fig. 15 [type locality: Ubay, Philippines, by present lectotype
designation, from Pinna sp.].
Type material. – Lectotype: SMF-ZMG956a (Go310b), female (10.6
× 8.7 mm), Burias, from Pinna sp. Paralectotypes: SMF-ZMG956b(Go310b), 3 females (9.5 × 7.5 – 10.5 × 9.0 mm), Burias, from
Pinna sp.; SMF-ZMG955 (Go310a), 2 females (7.2 × 6.3 – 8.7 ×
7.3 mm), Palau, from Pinna nigrina, coll. Semper.

Description. – Female: Carapace subhexagonal, wider thanlong, anterior margin well defined; front slightly projecting,transverse; posterior margin faintly convex. Eyes small,partially visible in dorsal view. MXP3 with propodus about
3 times as long as high, apex blunt, subtruncate; dactylus notreaching propodal apex; ischiomerus about twice as long as
wide, outer margin convex, inner margin with proximal 2/3
concave, angular at distal third; exopod outer margin
relatively straight, inner margin slightly convex, flagellum
two-segmented. Cheliped with dactylus about half palm
length; palm about twice as long as high, mesioventral marginsetose; dactylus occlusal margin with large proximal tooth;pollex occlusal margin irregular, sparsely setose. Walking
legs dorsally and ventrally unarmed; relative lengths of meri
P4 > P2 > P3 > P5. P2–P5 dactyli relative lengths P5 > P4
> P3 = P2. P2 and P3 dactyli about half propodal length. P4
asymmetrical, sparsely setose; longer dactylus shorter thanpropodus, about twice length of shorter dactylus; dactyli finely
dentate ventrally. P5 dactylus slightly longer than propodus;
ventrally setose, dorsally setose distally; with 2 rows of
subdistal spines, upper row with 7 or 8 spines, lower row

Fig. 5. Arcotheres latifrons (Bürger, 1895), new combination, USNM 32436, female lectotype (9.5 × 8.1 mm), Bohol, the Philippines: A,
dorsal habitus; B, anterior carapace; C, right cheliped; D, right MXP3. Scale bars: A = 2.0 mm; B, C = 1.0 mm; D = 0.7 mm.

198

THE RAFFLES BULLETIN OF ZOOLOGY 2007
with 16–18 spines, decreasing in size proximally. Abdomen
extending to buccal region, covering bases of walking legs.
Male: not known.
Remarks. – Six of the original eight syntypes of A. latus areextant – four from Burias, the Philippines, and two from
Palau. A female syntype (10.6 × 8.7 mm) from Burias,
Philippines, is herein designated as the lectotype of A. latus.

Arcotheres modiolicola (Bürger, 1895)(Fig. 7)
Pinnotheres modiolicola Bürger, 1895: 370, Pl. 9: Fig. 9, Pl. 10:
Fig. 9 [type locality: the Philippines, from Modiolaphilippinarium].
Type material. – Holotype: SMF-ZMG168 (Go951a), damaged
ovigerous female (8.0 × 6.5 mm), Philippines, from Modiolaphilippinarium, C. Semper.

Description. – Female: Carapace subhexagonal (incomplete
in holotype), slightly wider than long, anterior margin wellblunt; front slightly projecting, transverse; posterior margin
slightly concave. Eyes small, visible in dorsal view. MXP3(based on type account) with propodus about twice as longas high, apex blunt, subtruncate; dactylus not reaching
propodal apex; ischiomerus less than twice as long as wide,
outer margin convex, inner margin with proximal 2/3 concave,rounded at distal third; exopod inner and outer margin
relatively straight, flagellum two-segmented. Cheliped withdactylus slightly longer than half palm length; palm about
1.5 times as long as high, mesioventral margin setose; dactylus
occlusal margin with large proximal tooth; pollex occlusal
margin with irregularly dentate proximally, sparsely setose.
Walking legs dorsally and ventrally unarmed; relative lengthsof meri P2–P4 subequal > P5. P2–P5 dactyli relative lengths
P5 > P4 > P3 > P2. P2 and P3 dactyli about 2/3 propodal
length. P4 asymmetrical, sparsely setose; longer dactylus asabout 3/4 propodus length, about 1.1 times length of shorter
dactylus. P5 dactylus longer than propodus; ventrally setose,
dorsally setose distally; ventrodistal margin with 6 or 7 minutespines. Abdomen extending to buccal region, covering basesof walking legs. Male: not known.
Remarks. – Campos & Manning (2001) transferred P.
modiolicola to Arcotheres. Unfortunately, the holotype is inpoor condition, with carapace damaged and MXP3 missing.
Therefore, in the above diagnosis, features of the MXP3 arebased on the type description and figures.

Fig. 6. Arcotheres latus (Bürger, 1895), new combination, SMF-ZMG956a, female lectotype (10.6 × 8.7 mm), Burias, the Philippines: A,
dorsal habitus; B, right MXP3. Scale bars: A = 2.0 mm; B = 0.7 mm.

199

Ahyong & Ng: Pinnotherid types of Bürger (1895)
Arcotheres nudifrons (Bürger, 1895)(Fig. 8)
Pinnotheres nudifrons Bürger, 1895: 378, Pl. 9: Fig. 22, Pl. 10: Fig.
20 [type locality: Lapinig, Philippines, by present lectotype
designation].
Type material. – Lectotype: SMF-ZMG952 (Go535a), female (6.0
× 4.8 mm), Lapinig, the Philippines.

Description. – Female: Carapace subhexagonal, wider thanlong, anterior margin blunt, front slightly projecting,transverse; posterior margin transverse. Eyes not visible indorsal view. MXP3 with propodus about 3 times as long as
high, apex blunt, rounded; dactylus not reaching propodal
apex; ischiomerus less than twice as long as wide, outermargin convex, inner margin with proximal 3/4 concave,
bluntly angular at distal quarter; exopod inner and outermargin convex, flagellum 2-segmented. Cheliped withdactylus about 2/3 palm length; palm almost twice as longas high, mesioventral margin setose; dactylus occlusal margin
with large proximal tooth; pollex occlusal margin with low
proximal tooth, sparsely setose. Walking legs dorsally andventrally unarmed; relative lengths of meri P4 > P3 > P2 >P5. P2–P5 dactyli relative lengths P5 > P4 > P3 > P2. P2

dactyli about half propodal length. P3 dactyli exceeding half
propodal length. P4 sparsely setose; longer dactylus shorter
than propodus and about 1.5 times as long as shorter dactylus;
setose ventrally and dorsally. P5 dactylus as long propodus;
ventrally and dorsally setose; with row of about 8 ventral
subdistal spinules. Abdomen extending to buccal region,
covering bases of walking legs. Male: not known.
Remarks. – Campos & Manning (2001) transferred P.
nudifrons to Arcotheres. One of the original two syntypes of
A. nudifrons is extant, and is herein designated as thelectotype.

Arcotheres palaensis (Bürger, 1895)(Fig. 9)Pinnotheres palaensis Bürger, 1895: 372–373, Pl. 9: Fig. 12, Pl.
10: Fig. 12 [type locality: Palau, by present lectotype
designation].
Type material. – Lectotype: SMF-ZMG948a (Go534d), female (10.2
× 8.4 mm), Palau, C. Semper. Paralectotypes: SMF-ZMG948b(Go534d), 3 females (10.3 × 8.3 – 10.8 × 8.9 mm) (1 with bopyrid),
type locality; SMF-ZMG947 (Go534a), 1 female (10.0 × 7.0 mm)

Fig. 7. Arcotheres modiolicola (Bürger, 1895), SMF-ZMG168, female lectotype (8.0 × 6.5 mm), the Philippines: A, anterior carapace; B,
right cheliped; C, D, left and right P2; E, F, left and right P3; G, H, left and right P4; I, J, right and left P5; K, left MXP3 (after Bürger,
1895: pl. 10, fig 9). Scale bar: A–I = 1.0 mm.

200

THE RAFFLES BULLETIN OF ZOOLOGY 2007
(shrivelled), type locality; SMF-ZMG177 (Go534c), 4 females (9.5
× 7.4 – 11.8 × 8.3 mm) (shrivelled), Ubay, from Placuna sella, C.
Semper (= A. guinotae Campos, 2001).

Description. – Female: Carapace subhexagonal, wider thanlong, anterior margin well defined, sharp; front faintly
projecting, slightly convex; posterior margin slightly concave.
Eyes small, not visible in dorsal view. MXP3 with propodus
about 3 times as long as high, apex blunt, rounded; dactylus
not reaching propodal apex; ischiomerus less than twice aslong as wide, outer margin convex, inner margin withproximal 2/3 slightly concave, angular at distal third; exopod
inner and outer margin relatively straight, flagellum twosegmented. Cheliped with dactylus about half palm length;

palm about 2.5 times as long as high, mesioventral marginsetose; dactylus occlusal margin with large proximal tooth;pollex occlusal margin with 2 blunt proximal teeth, sparsely
setose. Walking legs dorsally and ventrally unarmed; relative
lengths of meri P3 > P2 = P4 > P5. P2–P5 dactyli relative
lengths: P5 > P4 > P3 > P2. P2 and P3 dactyli less than half
propodal length. P4 asymmetrical, sparsely setose; longer
dactylus slighty shorter than propodus, about 1.1 times lengthof shorter dactylus. P5 dactylus slightly longer than propodus;
ventrally setose; dorsally setose distally. Abdomen extending
to buccal region, covering bases of walking legs.
Remarks. – All the syntypes of A. palaensis are extant except
for the specimens from Burias, the Philippines (two males

Fig. 8. Arcotheres nudifrons (Bürger, 1895), SMF-ZMG952, female lectotype (6.0 × 4.8 mm), Lapinig, the Philippines: A, dorsal habitus;B, right cheliped; C, right MXP3. Scale bars: A = 2.0 mm, B, C = 1.0 mm.

201

Ahyong & Ng: Pinnotherid types of Bürger (1895)
and two females). The remaining syntype series of A.
palaensis, however, includes two species, one species from
Palau and another from Ubay, the Philippines. Notably, thosefrom Ubay are referable to A. guinotae Campos, 2001.
Therefore, a syntype from Palau is designated as the lectotypeof the species to fix the identity of A. palaensis in line with
its specific epithet and to preserve the name A. guinotae as
defined by Campos (2001). Campos & Manning (2001)
attributed a geographic range to A. palaensis that spans Palau,
the Philippines, Indonesia and Malaysia. With the present
recognition of heterogeneity in the type series and fixationof a lectotype for A. palaensis, all records of the species
outside Palau require verification.

Arcotheres pernicola (Bürger, 1895), new combination(Fig. 10)
Pinnotheres pernicola Bürger, 1895: 375–376, Pl. 9: Fig. 17, Pl.
10: Fig. 16 [type locality: Ubay, the Philippines, from Perna,
by present lectotype designation].
Type material. – Lectotype: SMF-ZMG178 (Go947a), ovigerous
female (6.0 × 5.0 mm), Ubay, from Perna, C. Semper, 1863–1864.

Diagnosis. – Female: Carapace broader than long; front not
protruding anteriorly beyond dorsal outline. MXP3 propodus
distally tapering; dactylus reaching propodal apex. Dactylus

Fig. 9. Arcotheres palaensis (Bürger, 1895), SMF-ZMG948a, female lectotype (10.2 × 8.4 mm), Palau: A, dorsal habitus; B, right cheliped;
C, right MXP3. Scale bars: A = 2.0 mm; B = 1.0 mm; C = 0.7 mm.

202THE RAFFLES BULLETIN OF ZOOLOGY 2007of cheliped about 2/3 as long as palm. P2–3 dactyli subequalin length, about half as long as respective propodi. P4
distinctly longer than other pereopods; dactylus about twice
length of P5 dactylus. P5 dactylus more than twice as longas those of P2–3. Male: not known.
Remarks. – The single extant syntype of A. pernicola is in
extremely poor condition: the carapace is shrivelled andcollapsed, most pereopods are broken or lost, and the left
and right MXP3 is missing. Thus, the syntype of A. pernicola
is effectively useless, and it would be futile to make alectotype designation. Should a specimen be found that bears
the above diagnostic characters, a case should be made to
the International Commission for Zoological Nomenclature
to set aside the syntype to permit the selection of a neotype.
We tentatively assign P. pernicola to Arcotheres on the basisof the elongate dactyli of P4 and P5, which are longer thanthose of P2–3. The remnants of the syntype show that the P5
dactylus is more than twice as long as those of P2–3. Bürger’s(1895) figure is inaccurate in showing the dactyli of P2–3
longer than half the length of the P5 dactylus.

Arcotheres rayi, new species(Fig. 11)
Xenophthalmus latifrons Bürger, 1895: 387 [part, not X. latifrons
Bürger, 1895].
Type material. – Holotype: SMF-ZMG952, female (11.3 × 8.7 mm),
Mariveles or Bohol, Semper (paralectotype of X. latifrons Bürger,1895).

Description. – Female: Carapace subtrapezoid, wider thanlong, anterior margin well defined; front not projecting but
demarcated by emargination above each orbit. Eyes not
visible in dorsal view. MXP3 with propodus greater than twice
as long as high, apex blunt; dactylus not reaching propodal
apex; ischiomerus about twice as long as wide, outer margin
convex, inner margin with proximal 3/4 slightly concave,angular at distal quarter; exopod inner and outer margin
convex, flagellum two-segmented. Cheliped with dactylus
about half palm length; palm about 3 times as long as high,mesioventral margin setose; dactylus occlusal margin withlarge proximal tooth; pollex occlusal margin with small
proximal tooth, sparsely setose. Walking legs dorsally andFig. 10. Arcotheres pernicola (Bürger, 1895), new combination, SMF-ZMG178, female syntype (6.0 × 5.0 mm), Ubay, the Philippines: A,
left cheliped; B, right P2; C, right P3; D, right P4 merus; E, left P5; F, dorsal habitus (after Bürger, 1895, Pl. 9: Fig. 17); G, left MXP3
(after Bürger, 1895, Pl. 10: Fig. 16). Scale bar = 1.0 mm.

203

Ahyong & Ng: Pinnotherid types of Bürger (1895)
ventrally unarmed; relative lengths of meri P4 > P3 > P2 >P5. P2–P5 dactyli relative lengths P4 > P5 > P3 = P2. P2 andP3 dactyli about half propodal length. P4 asymmetrical inlength; merus 2/3 carapace length; longer dactylus shorter
than propodus, about twice length of shorter dactylus. P5
dactylus shorter than propodus; ventrally setose. Abdomen
seven-segmented, extending to buccal region, covering basesof walking legs. Male: not known.
Etymology. – The species is named after our good friend, thelate Ray Manning, who helped and inspired us over many
years, and who both of us “blame” for getting us started on
this “most terrible nightmare” of pinnotherid systematics.

Remarks. – As had been discussed earlier, the type series of
Xenophthalmus latifrons contained two species, of which thesmaller syntype (9.5 × 8.1 mm, USNM 32436), was selected
above as the lectotype. The paralectotype (11.3 × 8.7 mm,
SMF-ZMG952) is here also made the holotype of a new
species, Arcotheres rayi. Arcotheres rayi closely resembles
A. latifrons in almost all respects, but is readily distinguishedby the following features: the presence rather than absenceof eyes, the longer P4 is proportionally longer than in A.
latifrons whereby the merus is two-thirds instead of half thecarapace length, the P5 dactylus is proportionally shorter thanrather than longer than the P4 dactylus, and the dactylus of
MXP3 does not reach to the end of the propodus.

Fig. 11. Arcotheres rayi, new species, SMF-ZMG952, female holotype (11.3 × 8.7 mm), Bohol, the Philippines (paralectotype of A. latifrons(Bürger)): A, dorsal habitus; B, anterior carapace; C, abdomen; D, right cheliped; E, right MXP3. Scale bars: A, C = 2.0 mm; B = 1.0 mm;
D, E = 0.7 mm.

204

THE RAFFLES BULLETIN OF ZOOLOGY 2007
A species that closely resembles A. latifrons and A. rayi is
the Indian species, Pinnotheres placunae Hornell &Southwell, 1909, sharing similar carapace morphology,
particularly in the flattened aspect and near straight anterior
margin bearing small orbital notches. The type descriptionof P. placunae is brief, and figures somewhat stylized,
depicting the walking legs as symmetrical left to right, with
subequal dactyli (contradicting the description in which theP4 and 5 dactyli are one and a half times the length of theP2 and 3 dactyli). A more detailed subsequent account of thespecies, however, indicates that the walking leg lengths areasymmetrical from left to right, particularly P4, and dactyl
lengths are as described by Hornell & Southwell (1909). As
such, P. placunae is referable to Arcotheres and is hereintransferred. Arcotheres placunae, new combination, like A.
rayi, is immediately distinguished from A. latifrons by thepresence of eyes. Arcotheres rayi and A. placunae are readily
distinguished by the relative lengths of the P4–5 dactyli:slightly exceeding twice the length of the P2–3 dactyli in theformer, and about one and a half times the length in the latter.

Arcotheres rhombifer (Bürger, 1895), new combination(Fig. 12)
Pinnotheres rhombifer Bürger, 1895: 374, Pl. 9: Fig. 15, Pl. 10:
Fig. 14 [type locality: Ubay, the Philippines, by present lectotype
designation, from Pectunculus aurifluus].

Fig. 12. Arcotheres rhombifer (Bürger, 1895), new combination, Ubay, the Philippines: A–C, ZMG179a, female lectotype (8.5 × 6.8 mm);
D, E, ZMG179b, female paralectotype (7.4 × 5.7 mm); A, dorsal habitus; B, E, right cheliped; C, MXP3; D, carapace. Scale bars: A, D =2.0 mm; B, E = 1.0 mm; C = 0.7 mm.

205

Ahyong & Ng: Pinnotherid types of Bürger (1895)
Pinnotheres latissimus Bürger, 1895: 373, Pl. 9: Fig. 13a, b, Pl. 10:
Fig. 13 [type locality: Manila, the Philippines] [new synonymy].
Type material of A. rhombifer. – Lectotype: SMF-ZMG179a(Go949a), ovigerous female (8.5 × 6.8 mm), Ubay, Philippines, from
Pectunculus aurifluus. Paralectotype: SMF-ZMG179b (Go949a), 1
ovigerous female (7.4 × 5.7 mm), type locality.

Description. – Female: Carapace subhexagonal, wider thanlong, anterior margin well defined; front faintly projecting,convex; posterior margin slightly concave. Eyes small, notvisible in dorsal view. MXP3 with propodus about 3 times
as long as high, tapering to blunt apex; dactylus not reaching
propodal apex; ischiomerus about twice as long as wide, outermargin convex, inner margin with proximal 2/3 straight,
angular at distal third; exopod inner margin relatively straight,
outer margin convex, flagellum two-segmented. Chelipedwith dactylus about half palm length; palm about 2.5 times
as long as high, mesioventral margin setose, ventral margin
with slight bulge near base of pollex; dactylus occlusal margin
with large proximal tooth; pollex occlusal margin with 2 blunt
proximal teeth, sparsely setose. Walking legs dorsally andventrally unarmed; relative lengths of meri P3 > P2 = P4 >P5. P2–P5 dactyli relative lengths P5 > P4 > P3 = P2. P2 andP3 dactyli less than half propodal length. P4 asymmetrical,
sparsely setose; longer dactylus as long as propodus, about
twice length of shorter dactylus. P5 dactylus longer thanpropodus; ventrally setose, dorsally setose distally. Abdomen
extending to buccal region, covering bases of walking legs.
Male: not known.
Remarks. – Campos & Manning (2001) transferred P.
rhombifer to Arcotheres. Both syntypes of A. rhombifer are
extant. The larger female syntype is herein designated as thelectotype.
The holotype of A. latissimus (type locality: Manila, thePhilippines) is lost, but comparison of the original accountand figures of the species with the types of A. rhombifer
suggests that the two species are conspecific. As figured by
Bürger (1895), the holotype of A. latissimus from Manila wasincomplete, having damaged P2 and P5, at least on the rightside. According to Bürger’s (1895) account and figures, A.
rhombifer is distinguished from A. latissimus by having a
proportionally longer carapace, with the width/length ratio
given as about 1.33 vs. 1.20, although based on Bürger’s
figures, the ratios are closer to 1.25 and 1.48 respectively.
According to measurements of the types of A. rhombifer, thewidth/length ratios are 1.28–1.30. Consequently, the carapace
ratios used by Bürger (1895) are neither reliable
distinguishing criterion, nor are they particularly accurate in
the case of A. latissimus. Both A. rhombifer and A. latissimus
are otherwise indistinguishable, sharing similar MXP3 andrelative lengths of the walking leg dactyli (where known).
Both names are clearly synonymous and as they were
published in the same paper, were select A. rhombifer as thesenior synonym of A. latissimus. To stablilise the identity ofthe two species, the lectotype of A. rhombifer is also
designated as the simultaneous neotype of A. latissimus. Thus,
A. latissimus becomes an objective junior synonym of A.
rhombifer.

206

Specimens from “Indo-Malayan Seas” first reported by Miers(1880) as “?Pinnotheres obesus” and subsequently identified
by Gordon (1936) as P. latissimus, resemble A. rhombifer,
notably in the slight bulge on the lower margin of the cheliped
palm. Gordon’s (1936) material, however, differs from A.
rhombifer in the dactylus of P5 being shorter rather thanlonger than that of P4, and as such probably represents anundescribed species.

Arcotheres rotundatus (Bürger, 1895)(Fig. 13)
Pinnotheres rotundatus Bürger, 1895: 378, Pl. 9: Fig. 21, Pl. 10:
Fig. 19 [type locality: Burias, the Philippines, by present
lectotype designation, from Circe].
Pinnotheres consors Bürger, 1895: 377, Pl. 9: Fig. 20, Pl. 10: Fig.
18 [type locality: Palau, from Circe] [new synonymy].
Type material of A. rotundatus. – Holotype: SMF-ZMG169(Go538a), ovigerous female (approx. 11.1 × 9.3 mm), carapace
shrivelled, Burias, from Circe, 1859–1864, C. Semper.

Description. – Female: Carapace wider than long, anterior
margin blunt, front projecting, relatively straight; posterior
margin convex. Eyes partially visible in dorsal view. MXP3
with propodus about 2.5 times as long as high, apex blunt,rounded; dactylus reaching to or slightly beyond propodal
apex; ischiomerus about 2.5 times as long as wide, outermargin convex, inner margin with proximal 3/4 concave,slightly angular at distal quarter; exopod outer margin
concave, inner margin relatively straight, flagellum twosegmented. Cheliped with dactylus about half palm length;
palm about twice as long as high, mesioventral margin setose;
dactylus occlusal margin with large proximal tooth; pollex
occlusal margin with 2 low proximal protrusions, sparsely
setose. Walking legs dorsally and ventrally unarmed; relative
lengths of meri P3 > P2 = P4 > P5. P2–P5 dactyli relative
lengths P5 = P4 > P3 > P2. P2 and P3 dactyli less than half
propodal length. P4 sparsely setose; longer dactylus shorter
than propodus and about twice as long as shorter dactylus,
with single subdistal ventral seta; shorter dactylus with row
of long setae on ventral margin. P5 dactylus slightly longer
than propodus; ventrally and laterally setose. Abdomen
extending to buccal region, covering bases of walking legs.
Male: not known.
Remarks. – Campos & Manning (2001) transferred both P.
rotundatus and P. consors to Arcotheres. The carapace ofthe holotype of A. rotundatus is partially crushed, but all
diagnostic features of the MXP3 and walking legs are intact.
Study of Bürger’s account indicates that A. rotundatus differsfrom A. consors (types lost) only in the length of the MXP3
dactyli, reaching the propodal apex in the latter and slightly
overreaching the apex in the former. As both species were
published in the same paper, we select Arcotheres rotundatus
as the senior synonym of A. consors. The holotype of A.
rotundatus is designated as the neotype of A. consors to fix
the identity of the latter as an objective junior synonym of
the former.

THE RAFFLES BULLETIN OF ZOOLOGY 2007
Arcotheres similis (Bürger, 1895)(Fig. 14)
Pinnotheres similis Bürger, 1895: 373–374, Pl. 9: Fig. 14 [typelocality: Ubay, the Philippines].
Type material. – Holotype: ZMG956 (Go950a), female (8.3 × 6.4mm), Ubay, the Philippines, C. Semper.

Description. – Female: Carapace subhexagonal, wider thanlong, anterior margin blunt; front faintly projecting,transverse; posterior margin slightly convex. Eyes small,visible in dorsal view. MXP3 with propodus about 3 times
as long as high, apex blunt, rounded; dactylus not reaching
propodal apex; ischiomerus about twice as long as wide, outermargin convex, inner margin with proximal 3/4 slightly
concave, angular at distal quarter; exopod inner margin
relatively straight, outer margin broadly convex, flagellum
two-segmented. Cheliped with dactylus almost 3/4 palm
length; palm about twice as long as high; dactylus occlusal
margin with large proximal tooth; pollex occlusal margin withlow blunt proximal teeth, sparsely setose. Walking legs
dorsally and ventrally unarmed; relative lengths of meri P4> P3 > P2 > P5. P2–P5 dactyli relative lengths P5 > P4 > P3
> P2. P2 and P3 dactyli longer than than half propodal length.
P4 asymmetrical; longer dactylus shorter than propodus, about
1.2 times length of shorter dactylus; ventral setae longer thandorsal setae, apex spiniform. P5 dactylus longer than propodus
and twice as long as P2 dactylus, 1.6 times as long as P3
dactylus; ventral setae longer than dorsal setae, apex
spiniform; with with 2 rows of subdistal spines, upper row
with 5 spines, lower row with 8–11 spines, decreasing in size
proximally. Abdomen extending to buccal region, coveringbases of walking legs. Male: not known.
Remarks. – The holotype of A. similis is extant and in goodcondition. Gordon (1936) described and figured a specimen
from Singapore identified as A. similis. Based on her account,however, the Singaporean specimen differs from the holotype
in having a more tapering propodal apex on MXP3, in having
a less pronounced anterointernal angle on the ischiomerus of
MXP3, in having a relatively shorter dactylus of P3 (ratio of
P3 : P5 dactylus 1 : 2 vs. 1 : 1.6) and in the presence of 8
short spinules rather than 5 or 6 slender spinules on the ventral
subdistal margin of the P5 dactylus. Gordon (1936) mentioned
only a single row of subdistal dactylar spines in her accountsof A. spinidactylus and A. similis, rather than two rows asobserved in the holotype of the latter. The lower row ofdactylar spines, however, can be obscured by setae and wasprobably overlooked by Gordon (1936). Arcotheres similis
otherwise agrees with A. spinidactylus (Gordon, 1936)
indicating that the two species are probably conspecific. Thus,
Gordon’s (1936) Singaporean specimen identified as A.

Fig. 13. Arcotheres rotundatus (Bürger, 1895), SMF-ZMG169, female holotype (approx. 11.1 × 9.3 mm), Burias, the Philippines: A, carapace;
B, C, left and right P2; D, left P3; E, F, left and right P4; G, left P5; H, right cheliped; I, right MXP3. Scale bars: A–H = 2.0 mm; I = 0.7
mm.

207

Ahyong & Ng: Pinnotherid types of Bürger (1895)
similis should be restudied; it probably represents anundescribed species.

Arcotheres sp.
Material. – ZMG 956, 10 females (9.0 × 6.8 – 9.7 × 7.6 mm), thePhilippines, C. Semper 1859–1864.

Remarks. – These 10 Arcotheres specimens were not reported
on by Nauck, Semper or Bürger. The specimens are in a poor
state of preservation and not identifiable to species, with legs,
mouthparts etc. missing or broken. Owing to the dimensionsof the specimens, however, it is clear that they do not represent
any of the missing types. Bürger may have excluded thismaterial from his report because of their incomplete condition.

Buergeres holothuriae (Semper, 1880)(Fig. 15)
Pinnotheres holothuriae Semper, 1880: 105 [type locality:
Zamboanga, the Philippines, by lectotype designation (Ng &Manning, 2003)]; Bürger, 1895: 381–382, Pl. 9: Fig. 27, Pl. 10:
Figs. 26, 36.
Buergeres holothuriae – Ng & Manning, 2003: 912, Fig. 5.
Type material. – Lectotype: USNM 32433a, ovigerous female (10.1
× 9.3 mm), Zamboanga, Philippines, Semper, 1876. Paralectotype:
USNM 032433b, 1 ovigerous female (8.0 × 6.5 mm), type locality.

Diagnosis. – Female: Carapace almost circular, slightly wider
than long, width length ratio 1.1–1.2; dorsal surface smooth,
gently convex. Lateral margins gently convex; sub-branchialsurfaces gently setose. MXP3 with ischium and merus

Fig. 14. Arcotheres similis (Bürger, 1895), ZMG956, female holotype (8.3 × 6.4 mm), Ubay, the Philippines: A, dorsal habitus; B, rightcheliped; C, right MXP3. Scale bars: A = 2.0 mm; B = 1.0 mm; C = 0.5 mm.

208THE RAFFLES BULLETIN OF ZOOLOGY 2007

Fig. 15. Buergeres holothuriae (Semper, 1880), USNM 32433a, female lectotype (10.1 × 9.3 mm), Zamboanga, the Philippines: A, dorsal
habitus; B, right MXP3 (setae omitted); C, left chela; D, right P4; E, right P5. Scale bars: A = 2.0 mm; B = 0.5 mm; C–E = 1.0 mm. (Modified
after Ng & Manning, 2003).

Fig. 16. Buergeres ortmanni (Bürger, 1895), ZRC 2003.59, female neotype (11.4 × 9.5 mm), Pulau Salu, northeastern Singapore: A, dorsal
habitus; B, left MXP3 (setae omitted); C, right cheliped; D–G, right P2–P5. Scale bars: A, C–G = 2.5 mm; B = 1.0 mm (Modified after
Ng & Manning, 2003).

209

Ahyong & Ng: Pinnotherid types of Bürger (1895)
completely fused, without trace of suture, subcircular, inner
margin rounded; exopod relatively stout, outer margin gently
convex, with well developed flagellal setae; palp threesegmented, propodus elongate, longer than dactylus; dactylus
spatuliform, articulation point slightly anterior to base of
propodus. Chelipeds subequal; inner surfaces of merus,
carpus, palm and fingers covered with relatively long setae;
outer surface of chela densely covered with short pubescence;palm relatively stout; fingers shorter than palm, basal third
covered with dense pubescence; cutting edges of dactylus andpropodus each with numerous denticles except for almost
unarmed distal third. Walking legs relatively slender, P3 >P2 > P4 > P5 in length; dactyli of P2–4 relatively short,distinctly hooked; dactylus of P5 very long, about 5 times aslong as those of other legs; dorsal and ventral margins ofmeri of all legs lined with setae of various lengths; ventral
margins of propodus and dactylus of all legs lined with setae.
Abdomen with all segments distinct, freely articulating, telson
semicircular. Male: not known.
Remarks. – Ng & Manning (2003) redescribed this species
and selected a lectotype.

Buergeres ortmanni (Bürger, 1895)(Fig. 16)
Pinnotheres ortmanni Bürger, 1895: 384–385, Pl. 9: Fig. 30, Pl. 10:
Fig. 28 [type locality: Pulau Salu, Singapore, by neotype
designation (Ng & Manning, 2003)].
Buergeres ortmanni – Ng & Manning, 2003: 911, Fig. 4.
Type material. – Neotype: ZRC 2003.59, female (11.4 × 9.5 mm),
Pulau Salu, northeast Singapore, in holothurian in coral reef, exposedat edge of reef, R. U. Gooding, 28 Apr.1967.

Diagnosis. – Female: Carapace almost circular, slightly wider
than long, width to length ratio 1.1–1.2; dorsal surface smooth,
gently convex; intestinal region slightly depressed. Lateral
margins gently convex; sub-branchial surfaces densely setose.
MXP3 with ischium and merus completely fused, without
trace of suture, subovate, inner margin subangular at distal
third; exopod relatively stout, outer margin gently convex,
with well developed flagellal setae; palp three-segmented,
propodus elongate, longer than dactylus; dactylus spatuliform,
articulation point near midlength of propodus. Chelipeds
subequal; inner surfaces of merus, carpus, palm and fingers
densely covered with relatively long setae; palm relatively
stout; fingers shorter than palm; cutting edges of dactylus
and propodus each with sub–basal large tooth and 1 or 2
denticles before it, rest of cutting edges smooth, almost blade–like; tips hooked. Walking legs relatively slender, P3 > P2
> P4 > P5 in length; dactyli of P2–4 relatively short, sharply
tapering; dactylus of P5 very long, about 3 times as long as
those of other legs; dorsal margins of meri of all legs relatively
densely lined with setae of various lengths, becoming sparseor absent along distal one-quarter; ventral margins ofpropodus of all legs relatively densely lined with setae on
distal half. Abdomen with all segments freely articulating,telson semicircular. Male: not known.

210

Remarks. – Ng & Manning (2003) redescribed this specieson the basis of a neotype from Singapore.

Buergeres tenuipes (Bürger, 1895)(Fig. 3B)
Pinnotheres tenuipes Bürger, 1895: 371–372, Pl. 9: Fig. 11, Pl. 10:
Fig. 11 [type locality: Ubay, Philippines, from holothurian].
Type material. – Lost.

Diagnosis. – Female: Carapace almost circular, slightly wider
than long; dorsal surface smooth, glabrous. MXP3 withischium and merus completely fused, inner margin obtusely
angled at distal third; palp three-segmented; dactylus
spatuliform, articulation point at midlength of propodus, apex
not overreaching propodus. Chelipeds inner surfaces ofmerus, carpus, palm and fingers densely covered withrelatively long setae; fingers shorter than palm; Walking legs
relatively slender; relative lengths: P3 > P2 > P4 > P5; dactyli
of P2–4 relatively short, sharply tapering; dactylus of P5 about
3 times as long as those of other legs; ventral margins ofmeri, propodi, and carpi of all legs setose. Male: not known.
[after Bürger, 1895: 371–372, Pl. 9: Fig. 11, Pl. 10: Fig. 11]
Remarks. – Pinnotheres tenuipes was assigned, with a bit of
doubt, to Buergeres by Ng & Manning (2003).

Durckheimia caeca Bürger, 1895
(Fig. 17)Durckheimia caeca Bürger, 1895: 385–386, Pl. 9: Fig. 33, Pl. 10:
Fig. 31 [type locality: Palau, from Lima squamosa Lamarck];– Ahyong & Ng, 2005: 119–121, Figs. 2, 3.
Type material. – Holotype: SMF-ZMG166, ovigerous female (10.2
× 9.0 mm), Palau, from Lima squamosa Lamarck, C. Semper.

Diagnosis. – Female: Carapace rounded in dorsal outline;
anterior margin rounded, with deep, U-shaped, median notch.
Median carina cristate with shallow concavity at about
midlength; extending from posterior margin to anterior thirdof carapace after which it descends to base of median anterior
notch. Endopod of MXP3 with ischiomerus about twice aslong as wide, inner margin slightly convex, outer margin
strongly convex; dactylus with apex slightly overreaching
propodal apex. Chela without dorsal carina on merus.
Pereopods 2–5 with merus cristate dorsally; carpus andpropodus rounded dorsally. Male: not known.
Remarks. – The species was redescribed by Ahyong & Ng
(2005).

THE RAFFLES BULLETIN OF ZOOLOGY 2007
Holotheres flavus (Nauck, 1880)(Fig. 18)
Pinnotheres flavus Nauck, 1880: 23, 66 [type locality: Bohol, Ubay,
the Philippines, by lectotype designation (Ng & Manning,2003)]; Bürger, 1895: 383–384, Pl. 9: Fig. 29, Pl. 10: Figs. 29,
35
Holotheres flavus – Ng & Manning, 2003: 907, Figs. 2, 3.
Type material. – Lectotype: SMF-ZMG 951a (Go297b), male (8.6
× 7.9 mm), Bohol, Ubay, the Philippines, C. Semper, 1863–1864.
Paralectotypes: SMF-ZMG 951b, 2 males (7.9 × 7.6 mm, 8.0 × 7.6mm), 2 females (10.3 × 10.0 mm, 8.9 × 8.0 mm), type locality;
USNM 32434, 2 ovigerous females (10.0 × 9.5 mm, 8.4 × 7.8 mm),
Zamboanga, the Philippines, C. Semper, 1876.

Diagnosis. – Carapace subcircular, width to length ratio ca.
1.1, margins setose, obscuring outline; anterolateral margin
arcuate, lateral margins straight, subparallel, posterolateral
margin gently convex. MXP3 with ischium and merus
completely fused, without trace of suture, subovate, inner
margin subangular submedially; exopod relatively stout, outermargin convex, with well developed flagellal setae; propodus
elongate, subconical, subequal in length to dactylus; dactylus
spatuliform, articulates at base of propodus. Chelipeds
subequal; inner surfaces of distal 4 segments covered withsetae; carpus dorsal surface entire, without dorsal groove;
palm relatively stout; fingers shorter than palm; cutting edgesof dactylus and propodus each with sub-basal large tooth andseveral denticles before it. Walking legs relatively stout, P3> P4 > P2 > P5 in length; dactyli of P2–5 relatively long,
sharply tapering, all subequal in length; dorsal and ventral
margins of meri, carpi and propodi of all legs lined with setaeof various lengths, obscures margins. Male and female
abdomens with all segments freely articulating.
Remarks. – Ng & Manning (2003) redescribed this species
and selected a lectotype.

Holotheres semperi (Bürger, 1895)(Fig. 19)
Pinnotheres semperi Bürger, 1895: 382–383, Pl. 9: Fig. 28, Pl. 10:
Fig. 27 [type locality: Singapore, by neotype designation (Ng& Manning, 2003)].
Holotheres semperi – Ng & Manning, 2003: 905, Fig. 1.
Type material. – Neotype: ZRC 1965.11.24.40, male (7.9 × 7.2 mm),in respiratory tree of Holothuria scabra, Singapore, S. H. Chuang,
1950s.

Diagnosis. – Female: Carapace of female subcircular, width
length ratio 1.1–1.2, anterolateral and lateral marginconfluent, gently convex; male carapace subquadrate, width
to length ratio ca. 1.1, anterolateral margin almost straight,
lateral margin gently convex; sub-branchial surface densely
setose dorsal surface gently convex, covered with dense layer

Fig. 17. Durckheimia caeca Bürger, 1895, SMF-ZMG166, holotype female (10.2 × 9.0 mm), Palau : A, dorsal habitus; B, right MXP3; C,carapace, right lateral view; D, carapace, anterior view; E, right P2. Scale bars: A, C = 3.0 mm; B = 0.7 mm; D, E = 1.5 mm. (Modified
after Ahyong & Ng, 2005, fig. 2).

211

Ahyong & Ng: Pinnotherid types of Bürger (1895)of short setae longer along lateral regions, completely
obscuring outline and surface; intestinal region slightly
depressed. MXP3 with ischium and merus completely fused,without trace of suture, subovate, inner margin obtusely
angular submedially; exopod relatively stout, outer margin
convex, with well developed setae on flagellum; propodus
elongate, conical, subequal in length to dactylus; dactylus
spatuliform, articulating at base of propodus. Chelipeds
subequal; surfaces of distal 4 segments densely covered withsetae, those on inner surfaces relatively longer; carpus dorsal
surface entire, without dorsal groove; palm relatively stout;
fingers shorter than palm; cutting edges of dactylus andpropodus each with sub-basal large tooth and numerous
denticles before and after it; distal-most part of fingers
excavated on inner surface. Walking legs relatively stout, P3> P4 > P2 > P5 in length; dactyli of P2–5 relatively long,
sharply tapering, all subequal in length; dorsal and ventral
margins of meri, carpi and propodi of all legs densely linedwith setae of various lengths, completely obscuring margins.
Male and female abdomens with all segments freely
articulating.
Remarks. – Ng & Manning (2003) recently redescribed thisspecies on the basis of a neotype from Singapore.

Fig. 18. Holotheres flavus (Nauck, 1880), Bohol, Ubay, the Philippines: A–H, SMF 951a, male lectotype (8.6 × 7.9 mm); I–K, USNM 32434,
female paralectotype (10.0 × 9.5 mm); A, I, dorsal habitus; B, J, right cheliped; C–F, left P2–P5; G, abdomen; H, left G1; K, left MXP3.
Scale bars: A–G, I, J = 1.0 mm; H = 0.3 mm; K = 2.0 mm. (Modified after Ng & Manning, 2003).

212

THE RAFFLES BULLETIN OF ZOOLOGY 2007
Holothuriophilus trapeziformis Nauck, 1880
(Fig. 20)

Nepinnotheres affinis (Bürger, 1895) new combination(Fig. 21)

Holothuriophilus trapeziformis Nauck, 1880: 24, 66 [type locality:
Mazatlan, Sinaloa, Mexico, by lectotype designation (Ng &Manning, 2003)]; Ng & Manning, 2003: 903, 916–918, Fig. 7C–F.
Pinnotheres trapeziformis – Bürger, 1895: 380–381, Pl. 9: Fig. 26,
Pl. 10: Fig. 25.

Pinnotheres affinis Bürger, 1895: 365–366, Pl. 9: Fig. 2, Pl. 10:
Fig. 2, 34 [type locality: Philippines, by present lectotype
designation, from Pinna].

Type material. – Lectotype: SMF-ZMG 170 (Go565a), female (7.7
× 4.8 mm), Mazatlan, Sinaloa, Mexico, from the cloaca of a
Holothuria inornata Semper.

Diagnosis. – Female: Carapace length about 0.6 times width,widest anterior to midlength. MXP3 palp three-segmented;
propodus conical distally, dactylus spatulate, broadening
distally, apex rounded, overreaching propodus, inserting near
midlength of propodus. Cheliped merus and carpus densely
setose; propodus setose on inner distal margin. Walking legs
similar, shorter than carapace width, segments compressed;
meri with setose dorsal and ventral margins; carpi and propodi
dactyli subequal. Abdomen seven-segmented.
Remarks. – Manning (1993b) resurrected Holothuriophilus
and Ng & Manning (1993) designated a lectotype for H.
trapeziformis.

Type material. – Lectotype: SMF-ZMG171a, ovigerous female (7.0
× 6.1 mm), the Philippines, from Pinna. Paralectotypes: ZMG171b,
1 male (6.7 × 6.3 mm), type locality; SMF-ZMG944 (Go539b), 7
males (2.5 × 2.5 – 5.7 × 5.3 mm), 3 females (5.6 × 4.9 mm – 8.4
× 7.6 mm), Bohol, C. Semper; USNM 32435, 1 female (9.5 × 8.2mm), Bohol, C. Semper. Probable paralectotypes: SMF-ZMG175(Go542a), 1 male (7.2 × 7.15 mm), 1 female (8.1 × 7.3 mm), Ubay,
C. Semper.

Description. – Female: Carapace subcircular, slightly wider
than long, front projecting slightly. Eyes small, not visible indorsal view. MXP3 with propodus about twice as long as
high, tapering distally to blunt apex; dactylus outreaching
propodal apex; ischiomerus outer margin convex, inner
margin with proximal 2/3 slightly concave; exopod inner andouter margin convex, flagellum two-segmented.
Cheliped with dactylus slightly longer than half palm length;
palm about twice as long as high, mesioventral margin setose;
dactylus occlusal margin finely dentate and sparsely setose,

Fig. 19. Holotheres semperi (Bürger, 1895). A–I, ZRC 1965.11.24.40, male neotype (7.9 × 7.2 mm). J, ZRC 1954.11.24.41, female (10.7
× 10.2 mm), Singapore: A, dorsal habitus (right side of carapace denuded); B, left MXP3; C, right chela; D–G, left P2–P5; H, abdomen;I, left G1; J, carapace; B–I, setae omitted. Scale bars: A–H = 1.0 mm; B; I = 0.5 mm; J = 1.5 mm. (Modified after Ng & Manning, 2003).

213

Ahyong & Ng: Pinnotherid types of Bürger (1895)
with large proximal tooth; pollex with low protrusion
proximally, margin sparsely setose; mesial margin of carpus
and mesiodistal margin of merus setose. Walking legs dorsallyand ventrally unarmed; relative lengths P3 > P2 = P4 > P5.
P2–P5 dactyli falcate; relative lengths P3 > P2 = P4 > P5.
P2–P5 propodus and carpus with ventral setae; P3–P5 merus
with dorsal setae. Abdomen extending to buccal region,
covering bases of walking legs. Male: Carapace subcircular,slightly wider than long, front not projecting. Eyes small,visible in dorsal view. MXP3 as in female. Cheliped withdactylus slightly longer than half palm length; palm length
less than twice as long as high, mesioventral margin setose;
dactylus occlusal margin finely dentate and sparsely setose,with large proximal tooth; pollex with low protrusion
proximally, margin sparsely setose; mesial margin of carpus
and mesiodistal margin of merus setose. Walking legs dorsallyand ventrally unarmed; relative lengths P3 > P2 = P4 > P5.
P2–P5 dactyli falcate; relative lengths P3 > P2 = P4 > P5.
P3–P5 propodus and carpus with ventral setae, merus withdorsal and ventral setae; P2 merus with ventral setae.
Abdomen widest at somite 3, tapering distally to evenly
rounded telson.
Remarks. – Pinnotheres affinis agrees in all respects with
Manning’s (1993) concept of Nepinnotheres and is hereintransferred. Most of the original syntypes of N. affinis, newcombination, are extant. The two syntypes from ZMG171
are illustrated; the female is herein designated as the lectotype.
One lot, ZMG175, containing one male and one female fromUbay, was accompanied by a recent label with the name
Pinnotheres gracilis. These specimens, however, are clearlyreferable to P. affinis rather than P. gracilis. Bürger listed
three specimens of N. affinis from Ubay, a male and two
females, as part of the original type series, so it is likely that
ZMG175 contains the remaining types of N. affinis from that
locality. As such, we treat them as probable paralectotypes.

Although Ng & Manning (2003) indicated that someparalectotypes of P. affinis remained in the Berlin Museum,
Oliver Coleman (pers. com. 2004) has informed us that none
are actually present in the Berlin collections.

Nepinnotheres cardii (Bürger, 1895) new combination(Fig. 22)
Pinnotheres cardii Bürger, 1895: 367–368, Pl. 9: Fig. 4, Pl. 10: Fig.
4 [type locality: Burias, the Philippines, by present lectotype
designation, from Cardium unedo].
Type material. – Lectotype: SMF-ZMG945 (Go955a), ovigerous
female (8.6 × 8.3 mm), Burias, from Cardium unedo, C. Semper,
1859–1864. Paralectotype: SMF-ZMG957 (Go955a), 1 male (4.4 ×
4.4 mm), type locality.

Description. – Female: Carapace subcircular, slightly wider
than long, front faintly projecting, slightly concave. Eyes
small, partially visible in dorsal view. MXP3 with propodus
more than twice as long as high, tapering distally to blunt
apex; dactylus not outreaching propodal apex; ischiomerus
about twice as long as wide, outer margin convex, inner
margin with proximal 2/3 slightly concave, bluntly angular
at distal third; exopod outer margin sinuous, inner margin
relatively straight, flagellum two-segmented. Cheliped withdactylus slightly longer than half palm-length; palm slightly
longer than twice as long as high, mesioventral margin setose;
dactylus occlusal margin finely dentate and sparsely setose,with large proximal tooth; pollex with low proximal tooth,margin sparsely setose; mesial margin of carpus setose.
Walking legs dorsally and ventrally unarmed; relative lengths
P3 > P2 > P4 > P5. P2–P5 dactyli falcate, sparsely setose;
relative lengths P2 = P 3 > P4 = P5. P4 and P5 propodus withsparse ventral setae. Abdomen extending to buccal region,
covering bases of walking legs. Male: Carapace subcircular,

Fig. 20. Holothuriophilus trapeziformis Nauck, 1880, SMF-ZMG 170, female lectotype (7.7 × 4.8 mm), Mazatlan, Sinaloa, Mexico: A,
dorsal habitus; B, right cheliped; C, right MXP3. Scale bars: A, B = 2.0 mm; C = 0.5 mm.

214

THE RAFFLES BULLETIN OF ZOOLOGY 2007

Fig. 21. Nepinnotheres affinis (Bürger, 1895), new combination, the Philippines: A–C, SMF-ZMG171a, female lectotype (7.0 × 6.1 mm);
D–F, SMF-ZMG171b, paralectotype male (6.7 × 6.3 mm); A, D, dorsal habitus; B, right cheliped; C, left MXP3; E, right MXP3; F, abdomen.
Scale bars: A, D, F = 2.0 mm; B = 1.0 mm; C, E = 0.7 mm.

215

Ahyong & Ng: Pinnotherid types of Bürger (1895)
as long as wide, front projecting, margin transverse. Eyes
small, visible in dorsal view. MXP3 as in female except
ischiomerus about 1.5 times as long as wide. Cheliped robust;
dactylus half palm length; palm height about 3/4 length;
dactylus occlusal margin with prominent, blunt proximal toothand lower distal tooth, sparsely setose; pollex occlusal margin
with irregular protrusions and few setae; mesial margin of
carpus setose. Walking legs dorsally and ventrally unarmed;

relative lengths P3 > P2 > P4 > P5. P2–P5 dactyli falcate,sparsely setose; relative lengths P3 > P2 > P4 > P5. P3–P5
propodus and carpus with dorsal and ventral setae. P4 andP5 merus with dorsal and ventral setae. Abdomen widest at
somite 3, tapering distally to evenly rounded telson.
Remarks. – Characters of P. cardii correspond well to
Nepinnotheres, to which the species is herein transferred. BothFig. 22. Nepinnotheres cardii (Bürger, 1895), new combination, Burias, the Philippines: A–C, SMF-ZMG945, female lectotype (8.6 × 8.3mm); D–G, SMF-ZMG957, paralectotype male (4.4 × 4.4 mm); A, D, dorsal habitus; B, E, right cheliped; C, right MXP3; F, left MXP3;
G, abdomen. Scale bars: A = 2.0 mm; B, D, E, G = 1.0 mm; C = 0.7 mm; F = 0.5 mm.

216

THE RAFFLES BULLETIN OF ZOOLOGY 2007
syntypes of N. cardii are extant. The right MXP3 of N. cardii
figured by Bürger (1895: pl. 10, fig 4) is from the male
syntype. The female syntype of N. cardii is here designatedthe lectotype of the species.

Remarks. – Characters of P. glaberrimus correspond well to
Nepinnotheres, to which the species is transferred. Two of
three of Bürger’s syntypes of N. glaberrimus are extant. The
lost syntype, from Ubay, the Philippines, was hosted by Lima
divaricata. The syntype from Zamboanga (SMF-ZMG 174)is herein designated the lectotype of the species.

Nepinnotheres glaberrimus (Bürger, 1895) new
combination(Fig. 23)
Pinnotheres glaberrimus Bürger, 1895: 366–367, Pl. 9: Fig. 3, Pl.
10: Fig. 3 [type locality: Zamboanga, the Philippines, by present
lectotype designation].
Type material. – Lectotype: SMF-ZMG174 (Go543a), male (5.3 ×
5.2 mm), Zamboanga, the Philippines, brackish water, C. Semper,
1859–1864. Paralectotype: SMF-ZMG185, 1 male (2.2 × 2.1 mm),
Palau, from Arca sp., C. Semper.

Description. – Female: not known. Male: Carapace
subcircular, slightly wider than long, front projecting, margintransverse. Eyes small, visible in dorsal view. MXP3 withpropodus more than twice as long as high, tapering distallyto blunt apex; dactylus not reaching propodal apex;
ischiomerus outer margin convex, inner margin with proximal
2/3 concave, margin at distal third produced to distinct angle;
exopod inner and outer margin convex, flagellum twosegmented. Cheliped robust; dactylus about 3/4 palm length;
palm length height about 2/3 length, mesioventral marginsetose; dactylus occlusal margin sparsely setose, with large
proximal tooth; pollex with 2 large proximal teeth, margin
sparsely setose; mesiodistal margin of merus setose. Walking
legs dorsally and ventrally unarmed; relative lengths P4 >P3 > P2 > P5. P2–P5 dactyli falcate, subequal. P2 carpus
with distal setae. P3 and P4 propodus and carpus with dorsal
setae. Abdomen widest at somites 2 and 3, tapering distallyto evenly rounded telson.

Nepinnotheres pectinicola (Bürger, 1895)
new combination(Fig. 3C)
Pinnotheres pectinicola Bürger, 1895: 362, 365, Pl. 9: Fig. 1, Pl.
10: Fig. 1 [type locality: Ubay, the Philippines, from Pecten
radula].
Type material. – Lost.

Diagnosis. – Female: Carapace subquadrate; front projecting
slightly beyond dorsal outline. Dactylus of MXP3
overreaching apex of propodus; propodus tapering to rounded
apex; ischiomerus with rounded innerdistal margin. Walking
legs with relative lengths P3 > P2 > P4 > P5; dactyli lessthan half propodus length. Male: not known. [asfter Bürger,
1895: 362, 365, Pl. 9: Fig. 1, Pl. 10: Fig. 1]
Remarks. – Pinnotheres pectinicola is tentatively assigned
to Nepinnotheres. Although species placed in Nepinnotheresby Manning (1993a) have a subcircular instead subquadrate
carapace, Bürger’s species otherwise appears to conform toManning’s original generic concept. Nepinnotheres
pectinicola might eventually have to be transferred to a new
genus when the genus is revised.
The holotype of N. pectinicola is lost, and at present, we do
not have material identifiable with the species. According to
the type account, the combination of the subquadrate carapace

Fig. 23. Nepinnotheres glaberrimus (Bürger, 1895), new combination, SMF-ZMG174, male lectotype (5.3 × 5.2 mm), Zamboanga, thePhilippines: A, dorsal habitus; B, right cheliped; C, left MXP3; D, abdomen. Scale bars: A, D = 2.0 mm; B = 0.7 mm; C = 0.5 mm.

217

Ahyong & Ng: Pinnotherid types of Bürger (1895)
and MXP3 dactylus over-reaching the propodal apex isdiagnostic for N. pectinicola.

Nepinnotheres rathbunae (Schmitt, McCain &Davidson, 1973) new combination(Fig. 3D)

Type material of P. glaber. – Lectotype: USNM 32431, ovigerous
female (7.8 × 6.5 mm), Palau, from Tapes turgida, C. Semper.
Paralectotypes: SMF-ZMG 173 (Go304a), 6 males (3.4 × 3.1 – 4.8
× 4.3 mm), 4 females (3.5 × 3.5 – 8.3 × 7.4 mm), type locality; type
locality.
Type material of P. impressus. – Syntypes: SMF-ZMG 176(Go953a), 2 females (6.0 × 4.8 – 6.1 × 4.5 mm), Aibukit, Palau, C.
Semper.

Pinnotheres barbatus Bürger, 1895: 369–370, Pl. 9: Fig. 8, Pl. 10:
Fig. 8 [type locality: Aibukit, Palau, from Donax sp.].
Orthotheres rathbunae Schmitt, McCain & Davidson, 1973: 27[replacement name for Pinnotheres barbatus Bürger, 1895,
preoccupied by Pinnotheres barbatus Desbonne, 1867].
Type material. – Lost.
Diagnosis. – Carapace circular, front slightly projecting beyond
margin, transverse. Dactylus of MXP3 articulating near proximal
third of ventral propodal margin, not reaching apex of propodus;
propodus height exceeding half length, apex broad, blunt;
ischiomerus with obtuse innerdistal angle. Cheliped and lower
margins of walkings legs densely setose. [after Bürger, 1895: 369–
370, Pl. 9: Fig. 8, Pl. 10: Fig. 8]

Remarks. – Schmitt et al. (1973) recognised that Pinnotheres
barbatus Bürger, 1895, was preoccupied by P. barbatus
Desbonne, 1867, and provided a replacement name,rathbunae for the former. Sakai (1969) placed P. barbatus
in Orthotheres despite the fact that the dactylus of the MXP3
is not distally articulated. Rather, characters of the MXP3
and dactyls of the walking legs of P. barbatus according to
Bürger (1895) correspond well with the genus Nepinnotheres,to which the species is transferred. The female holotype of
N. barbatus is lost.

Nepinnotheres villosulus (Guérin-Méneville, 1831) new
combination
Pinnotheres villosulus Guérin-Méneville, 1831: 7, Pl. 4: Fig. 6 [typelocality: Timor, Indonesia].
Pinnotheres villosus [sic] – Bürger, 1895: 366, Pl. 10: Fig. 5.
Material. – Lost.

Remarks. – Under the mis-spelt name “Pinnotheres villosus”,
Bürger (1895) reported eight specimens from two Philippine
localities: Ubay and Zamboanga. None appear to be extant.
Characters of P. villosulus, particularly the subequal walking
leg dactyli and articulation of the MXP3 dactylus near the
midlength of the propodus indicate that the species belongs
in Nepinnotheres. Hosts of N. villosulus reported by Bürger
(1895) are Pinna chemnitzii and Meleagrina margaritifera.

Orthotheres glaber (Bürger, 1895) new combination(Fig. 24, Tab. 1)
Pinnotheres glaber Bürger, 1895: 379, Pl. 9: Fig. 23, Pl. 10: Fig.
21 [type locality: Palau, by present lectotype designation, from
Tapes turgida].
Pinnotheres impressus Bürger, 1895: 380, Pl. 9: Fig. 24, Pl. 10:
Fig. 23 [type locality: Aibukit, Palau] [new synonymy].

218

Description. – Female: Carapace subcircular, slightly wider
than long, front faintly projecting, convex. Eyes small, notvisible in dorsal view. MXP3 with propodus more than twice
as long as high, apex rounded; dactylus inserting subdistally,outreaching propodal apex; ischiomerus exceeding twice
length, outer margin convex, inner margin with proximal 3/
4 concave, angular at distal quarter; exopod outer margin
convex, inner margin relatively straight, flagellum twosegmented. Cheliped with dactylus about 2/3 palm length;
palm 2.5 times as long as high, mesioventral margin setose;
dactylus occlusal margin finely setose, with large proximal
tooth; pollex occlusal margin sinuous, sparsely setose.
Walking legs asymmetrical in length from right to left;
propodi broadening distally; relative lengths P3 > P4 > P2 >P5. P2–P5 dactyli sparsely setose, shorter than half-length ofrespective propodi; relative lengths P3 > P4 > P2 > P5. P4
and P5 propodus with sparse ventral setae. Abdomen
extending to buccal region and covering bases of walking
legs. Male: Carapace subquadrate, wider than long, front
projecting, medially emarginate. Eyes visible in dorsal view.
MXP3 as in female, except with small distinct tooth at innerdistal angle of ischiomerus. Cheliped robust; dactylus almost
as long as palm; palm almost as high as long, outer surface
with shallow, irregular pits; dactylus occlusal margin withprominent, blunt proximal tooth, distal margin crenulate,sparsely setose; pollex occlusal margin finely dentate, sparsely
setose; mesial margin of carpus setose. Walking legs dorsallyand ventrally unarmed, length asymmetrical from left to right;
relative lengths P3 > P2 > P4 > P5. Dactyli shorter than halflength of respective propodi; P2–P4 dactyli subequal in lengthand slightly longer than P5 dactylus. P3 and P4 propodus
with dorsal and ventral setae; carpus with diagonal setal row.
Abdomen widest at somites 3 and 4, tapering distally to evenly
rounded telson.
Remarks. – Pinnotheres glaber is transferred to Orthothereson the basis of the subdistal insertion of the dactylus on the
propodus of the MXP3. Ng & Manning (2003) indicatedspecimens of O. glaber are present in the Berlin Museum,but since all 11 specimens listed by Bürger (1895) areaccounted for here, the Berlin specimens are not part of thetype series. The ovigerous female specimen (USNM 32431)
is herein selected as the lectotype. Bürger (1895) distinguishedO. glaber from a very similar species, O. impressus, also
described from Palau, by the relative lengths of the P2 andP5, with P2 subequal in length to P5 in the former, and P2
distinctly longer than P5 in the latter. Comparison of the intact
female types of O. glaber, however, indicates that Bürger’s(1895) diagnostic characters are inadequate. The walking legsin O. glaber are asymmetrical from left to right. In thelectotype, the ratio of P2 length to P5 length on the left andTHE RAFFLES BULLETIN OF ZOOLOGY 2007
right sides is 1.1 and 1.3 respectively. Similarly, the relative
lengths of P2 and P5 varied from side to side in the four otherfemale paralectotypes for which the requisite limbs are intact
on both sides (Table 1). Thus, the specimens could be
identified either as O. glaber or O. impressus depending on
the side for which P2 and P5 are compared. Evidently, Bürger
(1895) described O. glaber and O. impressus without
considering the pereopodal asymmetry. Thus, O. impressus
herein synonymised with O. glaber. As both names were
published in the same paper, we here choose Pinnotheres

glaber Bürger, 1895, as having priority over Pinnotheres
impressus Bürger, 1895, when the two names are considered
to be synonymous.
Remains of both original syntypes of O. impressus are extant,but in extremely poor condition: all limbs are broken andincomplete, left and right MXP3 are missing, and carapace
remains are soft and disintegrating. It seems pointless to selecta lectotype from this series.

Fig. 24. Orthotheres glaber (Bürger, 1895), new combination, Palau: A–C, USNM 32431, female lectotype (7.8 × 6.5 mm); D–G, ZMG
173, male paralectotype (4.0 × 3.6 mm); A, D, dorsal habitus; B, E, right cheliped; C, F, right MXP3; G, abdomen. Scale bars: A = 2.0 mm;
B, D, E, G = 1.0 mm; C = 0.4 mm; F = 0.2 mm.

219

Ahyong & Ng: Pinnotherid types of Bürger (1895)
Table 1. Relative lengths of P2 and P5 for left and right sides of female Orthotheres glaber.
Specimen

P2: P5 length (left)

P2: P5 length (right)

Paralectotype, female 3.5 × 3.5 mm
Paralectotype, female 5.2 × 4.5 mm
Paralectotype, female 7.5 × 6.3 mm
Lectotype, female 7.8 × 6.5 mm

1.3
1.4
1.2
1.1

1.7
1.2
1.4
1.3

Orthotheres laevis (Bürger, 1895)(Fig. 3E)

Paralectotypes: SMF-ZMG946b (Go542a), 2 females (5.6 × 4.7–
6.3 × 5.7 mm), type locality.

Pinnotheres laevis Bürger, 1895: 380, Pl. 9: Fig. 25, Pl. 10: Fig. 24[type locality: Palau, from Coralliophaga sp.].

Description. – Female: Carapace subcircular, slightly wider
than long, front faintly projecting, slightly concave. Eyes
small, partially visible in dorsal view. MXP3 propodus about
3 times as long as high, margins subparallel, apex blunt;
dactylus not reaching propodal apex; ischiomerus about twice
as long as wide, outer margin convex, inner margin withproximal 3/4 slightly concave, distinctly angular at distal
quarter; exopod inner and outer margin convex, flagellum
two-segmented. Cheliped with dactylus slightly longer thanhalf palm length; palm slightly shorter than twice as long as
high, mesioventral margin setose; dactylus occlusal margin
finely dentate, with large proximal tooth; pollex with occlusal
margin irregularly crenulate, sparsely setose; mesial marginof carpus setose. Walking legs dorsally and ventrally
unarmed; all segments with dorsal and ventral setate; relative
lengths P3 > P2 > P4 > P5. P2–P5 dactyli weakly falcate;
relative lengths P3 > P2 > P4 > P5. P2 dactylus 0.4–0.5
propodus length. P3 dactylus 0.4 propodus length. P4 dactylus
about 0.4 propodus length. P5 dactylus about 0.5 propodus
length. Abdomen extending to buccal region, covering basesof walking legs. Males: not known.

Type material. – Lost.

Diagnosis. – Carapace subhexagonal. MXP3 with dactylus
inserting subdistally, outreaching propodal apex; propodus
slightly longer than high; ischiomerus inner margin sinuous,
length almost 3 times width. Walking legs with propodi
subcylindrical, not inflated; P3 dactylus longer than P2, P4
and P5 dactyli. [after Bürger, 1895: 380, Pl. 9: Fig. 25, Pl.
10: Fig. 24]
Remarks. – The type account and figures indicate that the
dactylus of the MXP3 in P. laevis inserts subdistally on the
propodus, and is thus placed in Orthotheres. The holotype ofO. laevis is no longer extant.

Orthotheres longipes (Bürger, 1895) new combination(Fig. 3F)
Pinnotheres longipes Bürger, 1895: 379–380, Pl. 9: Fig. 31, Pl. 10:
Fig. 22 [type locality: Aibukit, Palau].
Type material. – Lost.

Diagnosis. – Female: Carapace subhexagonal. MXP3 withdactylus inserting subdistally, outreaching propodal apex;
propodus length about twice height; ischiomerus with obtuse
inner distal angle, length about twice width. Walking legswith propodi subcylindrical, not inflated; P3 dactylus longer
than P2, P4 and P5 dactyli. Male: not known. [after Bürger,
1895: 379–380, Pl. 9: Fig. 31, Pl. 10: Fig. 22]
Remarks. – The dactylus of the MXP3 in P. longipes inserts
subdistally on the propodus; the species is thus transferred toOrthotheres. The female holotype of O. longipes is no longer
extant.

Viridotheres gracilis (Bürger, 1895) new combination(Fig. 25)
Pinnotheres gracilis Bürger, 1895: 368–369, Pl. 9: Fig. 6, Pl. 10:
Fig. 6 [type locality: Ubay, the Philippines, by present lectotype
designation, from Solen].
Type material. – Lectotype: SMF-ZMG946a (Go542a), female (6.3
× 5.7 mm), Ubay, from Solen, C. Semper, 1862–1863.

220

Remarks. – Pinnotheres gracilis is herein transferred to
Viridotheres: it conforms well to Manning’s (1996) diagnosisof the genus, notably in the subcircular carapace, in P3 having
the longest dactylus, and being the longest of the walking
legs, and in having the dactylus of the MXP3 inserting near
the midlength on the ventral propodus margin. All three
original syntypes of V. gracilis are extant. The largest syntypeof V. gracilis is herein designated as the lectotype of thespecies.

Viridotheres otto, new species(Fig. 26)
Type material. – Holotype: ZMG 953a, ovigerous female (4.8 ×
3.7 mm), Philippines, C. Semper 1859–1864. Paratype: ZMG 953b,
1 immature female (2.8 × 2.3 mm), type locality.

Diagnosis. – Carapace front projecting slightly, transverse.
MXP3 propodus tapering distally to blunt apex. Cheliped
pollex with proximal tooth. Walking leg dactyli elongate,
apices falcate; relative lengths P3 > P2 > P4 > P5. P2 dactylus
about 0.5 times propodus length. P3 dactylus 0.5–0.6 times
propodus length. P4 dactylus about 0.6 times propodus length.
Description. – Female: Carapace subcircular, slightly wider
than long, front projecting slightly, transverse. Eyes small,THE RAFFLES BULLETIN OF ZOOLOGY 2007
visible in dorsal view. MXP3 with propodus about twice aslong as high, tapering distally to blunt apex; dactylus notreaching beyond propodal apex; ischiomerus outer margin
convex, inner margin with proximal 3/4 slightly concave;
inner distal angle blunt, obtuse; exopod inner margin straight,
outer margin convex, flagellum two-segmented. Chelipedwith dactylus about 2/3 palm length; palm less than twice aslong as high, mesioventral margin setose; dactylus occlusal
margin sparsely setose, with large proximal tooth; pollex withproximal tooth, margin sparsely setose; carpus and merus
glabrous. Walking legs dorsally and ventrally unarmed;
glabrous; relative lengths P3 > P2 = P4 > P5. P2–P5 dactyli
elongate, apices falcate; relative lengths P3 > P2 > P4 > P5.
P2 dactylus about 0.5 propodus length. P3 dactylus 0.5–0.6
propodus length. P4 dactylus about 0.6 propodus length. P5
dactylus about 0.6 propodus length. Abdomen sevensegmented, extending to buccal region, covering bases ofwalking legs. Male: not known.

Viridotheres was originally characterised by having P3 longer
than other walking legs, with the P3 dactylus distinctly longer
than those of other walking legs (Manning, 1993b). In V. otto,
new species, the dactylus of P3 is only slightly longer thanthat of P2. Although the new species perhaps belongs in a
separate genus, it is here referred to Viridotheres. The adultfemale holotype differs from the juvenile female paratype incarapace shape, being subcircular with a straight frontal
margin instead of being subtrapezoid with a concave frontal
margin. In other respects, the two specimens of V. otto agree
well. The new species most closely resembles Pinnotheres
burgeri Rathbun, 1909, and V. gracilis Bürger, 1895, in theP3 being longer than the remaining pereopods, and in thedactyli of P2–3 being longer than those of P4–5, with the P3
dactylus longer than the P2 dactylus. On the basis of therelative length of P3 and its dactylus in comparison to theremaining walking legs, Pinnotheres burgeri also belongs in
Viridotheres.

Etymology. – Named in honour of Otto Bürger, in recognitionof his major contributions to the study of the Pinnotheridae.
The name is used as a noun in apposition.

Viridotheres otto is distinguished from V. gracilis in having
glabrous instead of setose walking legs, in the tapering rather
than rounded propodal apex on MXP3, in the more strongly
curved dactylar apices of P2–4 dactyli, and in the longer P3–
4 dactyli (0.5–0.6 versus 0.4 propodus length). Viridotheres
otto differs from V. burgeri in the shorter dactyli on thewalking legs with the P2–5 dactyli about dactyli not exceeding

Remarks. – Although part of the Semper collection, these
specimens were not reported on by Bürger (1895). The
assignment of our new species to Viridotheres is tentative.

Fig. 25. Viridotheres gracilis (Bürger, 1895), new combination, SMF-ZMG946a, female lectotype (6.3 × 5.7 mm), Ubay, the Philippines:
A, dorsal habitus; B, right cheliped; C, right MXP3. Scale bars: A = 2.0 mm; B = 1.0 mm; C = 0.7 mm.

221

Ahyong & Ng: Pinnotherid types of Bürger (1895)
0.6 their respective propodal lengths instead of about 3/4
propodal length; in the shorter MXP3 dactylus, which falls
short of instead of reaching the propodal apex; and in the
narrower, more tapering MXP3 propodus. In addition, thedactylus of P3 in V. otto is only slightly longer than, insteadof distinctly longer than the P2 dactylus as in V. burgeri.
Unfortunately, the holotype of V. burgeri is a juvenile, so
the adult carapace shape is not known. The carapace shapeof the holotype of V. burgeri, however, is similar to that of
the juvenile paratype of V. otto, so both species possibly sharea similar adult carapace form.

Material. – ZMG167, 2 males (cw. 10.4–6.0 m, cl. 9.4–5.3 mm),
1 female (cw. 13.6 mm, cl. 11.9 mm), Bohol, coll. Semper.

Remarks. – The three specimens reported by Bürger (1895)
were reported by Ahyong & Ng (2005).

Xanthoidea sp.
Material. – ZMG 954, 13 juvenile males, the Philippines, C. Semper
1859–1864.

Remarks. – The specimens, collected by Semper from thePhilippines, and apparently unpublished, are very juvenile
xanthoids.

Xanthasia murigera White, 1846
Xanthasia murigera White, 1846: 177, Pl. 2, Fig. 3a [type locality:
Philippines]; Bürger, 1895: 386–387; Ahyong & Ng, 2005: 126–
127, Fig. 7.

Fig. 26. Viridotheres otto, new species, the Philippines: A–D, SMF-ZMG 953a, female holotype (4.8 × 3.7 mm); E, ZMG 953b, immature
female paratype (2.8 × 2.3 mm), carapace ; A, dorsal habitus ; B, right cheliped; C, right MXP3; D, abdomen; E, carapace. Scale bars: A,
D, E = 2.0 mm; B = 1.0 mm; C = 0.8 mm.

222

Extant

1 M, 1 F, the Philippines
4 F, 7 M, Bohol

223
2F extant

4 F, Palau

2 F, Lapinig
5 F, 6 M, Palau
1 F, Aibukit

Pinnotheres nudifrons Bürger, 1895

Pinnotheres glaber Bürger, 1895

Pinnotheres longipes Bürger, 1895

Lost

All extant; F lectotype designated1 F extant; lectotype designated

ExtantLost; neotype designated1 F, Palau
1 F, Burias

Pinnotheres consors Bürger, 1895

1 F extant; lectotype designatedLost

1 F, Ubay
4 F, Samar Island

Extant

1 F, Ubay, from Perna

Pinnotheres rotundatus Bürger, 1895

Pinnotheres exiguus Bürger, 1895

Pinnotheres pernicola Bürger, 1895

All extant; lectotype designated4 F, Burias

Pinnotheres latus Bürger, 1895

ExtantBoth extant; lectotype selected

1 F, Ubay
2 F, Ubay

Pinnotheres similis Bürger, 1895

Lost

Orthotheres longipes (Bürger, 1895)

Orthotheres glaber (Bürger, 1895)

Arcotheres nudifrons (Bürger, 1895)

Arcotheres rotundatus (Bürger, 1895)

Arcotheres rotundatus (Bürger, 1895)

Arcotheres exiguus (Bürger, 1895)

Arcotheres pernicola (Bürger, 1895)

Arcotheres latus (Bürger, 1895)

Arcotheres rhombifer (Bürger, 1895)

Arcotheres similis (Bürger, 1895)

Arcotheres rhombifer (Bürger, 1895)

A. guinotae Campos, 2001

ExtantLost

4 F, Ubay
2 M, 2F, Burias
1 F, Manila

A. palaensis (Bürger, 1895)

ExtantArcotheres palaensis (Bürger, 1895)

Buergeres tenuipes (Bürger, 1895)

Arcotheres arcophilus (Bürger, 1895)

4 F, Palau

Pinnotheres rhombifer Bürger, 1895

Pinnotheres latissimus Bürger, 1895

Nepinnotheres rathbuni Schmitt, McCain & Davidson, 1973
Arcotheres modiolicola (Bürger, 1895)

F, designated as lectotype

Lost

All extant; F lectotype selected

ExtantArcotheres coarctatus (Bürger, 1895)

Viridotheres gracilis (Bürger, 1895)

Nepinnotheres cardii (Bürger, 1895)

Nepinnotheres glaberrimus (Bürger, 1895)

Nepinnotheres affinis (Bürger, 1895)

Nepinnotheres pectinicola (Bürger, 1895)

Current name

1 F, Palau

1 F, Ubay

Pinnotheres tenuipes Bürger, 1895

Pinnotheres palaensis Bürger, 1895

2 F, 1M, Ubay

Pinnotheres arcophilus Bürger, 1895

Lost

1 F, Aibukit
1 F, the Philippines

Pinnotheres barbatus Bürger, 1895

Pinnotheres modiolicola Bürger, 1895

Lost

1 F, Zamboanga

Pinnotheres coarctatus Bürger, 1895

All extant; lectotype designated3 F, Ubay

Pinnotheres gracilis Bürger, 1895

Lost

1 M, Ubay
Both extant; F lectotype designated

Extant

1 M, Palau
1 M, 1 F Burias

Extant, lectotype designated

1 M, Zamboanga

Pinnotheres cardii Bürger, 1895

Pinnotheres glaberrimus Bürger, 1895

Extant, F lectotype designated

1 M, 2 F, Ubay

Lost
1M, 1F extant

1 F, Ubay

Pinnotheres pectinicola Bürger, 1895

Pinnotheres affinis Bürger, 1895

Current status

Material originally listed

Original name*

Table 2. Species reported by Bürger (1895) and unpublished specimens from the ‘Bürger collection’. Species listed in order of appearance in Bürger (1895). F = female; M = male

THE RAFFLES BULLETIN OF ZOOLOGY 2007

Lost
Lectotype female designated by
Ng & Manning (2003)

1 F, no data
1 F, Mazatlan

Holothuriophilus trapeziiformis Nauck, 1880

1 F, Mariveles or Bohol

Xenophthalmus latifrons Bürger, 1895

224
2 M, 1 F, Bohol
Not listed, Philippines
Not listed, Philippines
Not listed, Philippines

Xanthasia murigera White, 1847

Pinnotheres sp.

Pinnotheres sp.

13 M

10 F

2F, types of V. otto new species

All extant

Lost

Lost

1 F, Ubay, from Meleagrina
2 F, Burias, from Mytilus

Lost

1 F, Ubay
ExtantLost

1 F, Burias

Lost

3 F, Ubay

Lost

Holotype of A. rayi new species

F lectotype designated.

ExtantLost. F neotype designated by
Ng & Manning (2003)

1 F, Ubay, from Pinna

2 F, Zamboanga

Pinnotheres sp.

Pinnotheres parvulus Stimpson, 1858

Pinnotheres villosulus Guérin-Méneville, 1831
(as P. villosus)

1 F, Palau

Durckheimia caeca Bürger, 1895
1 F, Mariveles or Bohol

1 F, Abukit

3 M, 2 F, Ubay

Pinnotheres ortmanni Bürger, 1895

2 F extant
All extant; M lectotype designated by
Ng & Manning (2003)

6 M, 6 F, Zamboanga

Pinnotheres flavus Nauck, 1880

Lost. M neotype designated by
Ng & Manning (2003)

1 M, 1 F, Java

Pinnotheres semperi Bürger, 1895

2 F extant; lectotype designated by
Ng & Manning (2003)

8 F, 3 M, Zamboanga

Pinnotheres holothuriae Semper, 1880

Lost

1 F, Palao

Pinnotheres laevis Bürger, 1895

Extant2 F, Aibukit

Pinnotheres impressus Bürger, 1895

Table 2. Cont’d

Xanthoidea sp.

Arcotheres sp.

Viridotheres otto new species

Xanthasia murigera White, 1847

Arcotheres alcocki (Rathbun, 1909)

Nepinnotheres villosulus (Guérin-Méneville, 1831)

Arcotheres rayi new species

Arcotheres latifrons (Bürger, 1895)

Durckheimia caeca Bürger, 1895

Buergeres ortmanni (Bürger, 1895)

Holotheres flavus (Nauck, 1880)

Holotheres semperi (Bürger, 1895)

Buergeres holothuriae (Semper, 1880)

Holothuriophilus trapeziiformis Nauck, 1880

Orthotheres laevis (Bürger, 1895)

Orthotheres glaber (Bürger, 1895)

Ahyong & Ng: Pinnotherid types of Bürger (1895)

THE RAFFLES BULLETIN OF ZOOLOGY 2007
publication of plate 7 was cited as 1831?–1834, but the text
appeared in 1844. We follow this citation until an accurate year
is determined for the plates.]

ACKNOWLEDGMENTS
We are grateful to Michael Türkay and Andreas Allspach(SMF) for entrusting the Bürger material to us for study, andtheir tremendous help in understanding the collection. Support
to the first author from a Sydney Grammar School Fellowship,
two Research Fellowships from the Raffles Museum of
Biodiversity Research, National University of Singapore,
Biosecurity New Zealand (contract ZBS 200524) and theFoundation for Research, Science and Technology (BBBI063)
is gratefully acknowledged. Oliver Coleman (Berlin Museum)and Rafael Lemaitre (USNM) have also been very kind with
our many enquires about their pinnotherid material. Tohru
Naruse and an anonymous reviewer are thanked forconstructive criticism of the manuscript. Financial support
by research grants to the second author from the National
University of Singapore is also gratefully acknowledged.

Holthuis, L. B. & R. B. Manning, 1990. Crabs of the subfamily
Dorippinae MacLeay, 1838, from the Indo-West Pacific region(Crustacea: Decapoda: Dorippidae). Researches on Crustacea.
Special Number 3: i–iii, 1–151.
Hornell, J. & T. Southwell, 1909. Description of a new species of
Pinnoteres from Placuna placenta, with a note on the genus.
Report on the Marine Zoology of Okhamandal in Kittiawar, I:
99–103, pl. 1.
Man, J. G. de., 1887. Report on the podophthalmous Crustacea ofthe Mergui Archipelago, collected for the trustees of the IndianMuseum, Calcutta, by Dr. John Anderson, F.R.S.,
Superintendent of the Museum – Part II. Journal of the Linnean
Society, London, (Zoology), 22: 65–128.

LITERATURE CITED

Man, J. G. de., 1887–1888. Bericht über die von Herrn Dr. J. Brock
im indischen Archipel gesammelten Decapoden und
Stomatopoden. Archiv für Naturgeschichte, 53(I, 2): 215–288,
pls. 7–10 [issued 1887]; 53(I, 3): 216–600, pls. 11–22a [issued
1888] [fide Clark et al., 1990].

Ahyong, S. T. & P. K. L. Ng, 2005. Review of Durckheimia and
Xanthasia, with descriptions of two new genera (Crustacea:
Decapoda: Brachyura: Pinnotheridae). Journal of Crustacean
Biology, 25(1): 116–129.

Manning, R. B., 1993a. West African pinnotherid crabs subfamily
Pinnotherinae (Crustacea, Decapoda, Brachyura). Bulletin duMuséum national d’Histoire naturelle, Paris, (4)15A(1–4): 125–
177.

Bürger, O., 1895. Ein Beitrag zur kenntniss der Pinnotherinen.
Zoologische Jahrbücher, Abtheilung für Systematik, Geographie
und Biologie der Thiere, 8: 361–390, pls. 9, 10.
Campos, E., 2001. A new crab species of the genus Arcotheres
Manning, 1993, from Thailand (Crustacea, Brachyura,
Pinnothuridae). Zoosystema, 23(3): 493–497.
Campos, E. & R. B. Manning, 2001. Authorship and diagnosis of
the genus Arcotheres Manning, 1993 (Crustacea: Brachyura:
Pinnotheridae). Raffles Bulletin of Zoology, 49: 167–170.
Campos, E., V. Díaz, & J. A. Gamboa-Contreras, 1998. Notes on
distribution and taxonomy of five poorly known species of
pinnotherid crabs from the eastern Pacific (Crustacea:
Brachyura: Pinnotheridae). Proceedings of the Biological Societyof Washington, 111: 372–381.
Clark, P. F., Harrison, K., & S. Goodman, 1990. Decapoda andStomatopod by De Man: 1887 or 1888? Archives of Natural
History, 17(1): 79–80.
Dai, A. & S. Yang, 1991. Crabs of the China Seas. China Ocean
Press, Beiking. 608 pp., 74 pls.
Desbonne, I., 1867. Brachyoures. In: Desbonne, I. & A. Schramm(Eds.), Crustacés de la Guadeloupe d’après un manuscrit duDocteur Isis Desbonne comparé avec les échantillons decrustacés de sa collection et les dernières publications de MM.
Henri de Saussure et William Stimpson. Imprimerie duGouvernement, Basse Terre, pp. ii + 60.
Gordon, I., 1936. On a few Indo-Pacific species of Pinnotheres,with special reference to asymmetry of the walking legs. Journalof the Linnean Society of London, Zoology, 40: 163–180.
Guérin-Méneville, F. E., 1829–1844. Iconographie du règne animalde G. Cuvier, ou représentation d’après nature de l’une desespèces les plus remarquables, et souvent non figurèes, de chaquegenre d’animaux, avec un text descriptif mis au courant de lascience: Ouvrage pouvant servir d’atlas à tous les traités dezoologie. 450 plates in 45 livaisons. Crustacea: 36 pls., 48 p.
Paris. [According to Holthuis & Manning (1990: 139), thecorrect date of the Crustacean plates is unresolved. The year of

Manning, R. B., 1993b. Three genera removed from the synonymy
of Pinnotheres Bosc, 1802 (Brachyura: Pinnotheridae).
Proceedings of the Biological Society of Washington, 106: 523–
531.
Manning, R. B., 1996. Viridotheres marionae, a new genus andspecies of pinnotherid crab from West Africa (Crustacea:
Decapoda: Brachyura). Zoologische Mededelingen, 70(19): 271–
273.
Miers, E. J., 1880. On a collection of Crustacea from the Malaysian
region.–Part II. Telphusidea, Catametopa, and Oxystomata.
Annals and Magazine of Natural History, series 5, 5: 304–317,
pl. XIV.
Nauck, E., 1880. Das Kaugerust der Brachyuren (mit beschreibung
neur gattungen und arten, Z. T. von C. Semper). Zeitscrift
wissenschaften Zoologische, Leipzig, 34: 1–69, pl. 1.
Ng, P. K. L. & R. B. Manning, 2003. On two new genera of pea
crabs parasitic in holothurians (Crustacea: Decapoda: Brachyura:
Pinnotheridae) from the Indo-West Pacific, with notes on allied
genera. Proceedings of the Biological Society of Washington,
116(4): 901–919.
Rathbun, M. J., 1909. New crabs from the Gulf of Siam. Proceedingsof the Biological Society of Washington, 22: 107–114.
Schmitt, W. L., J. C. McCain, & E. Davidson, 1973. Decapoda I.
Brachyura I. Fam. Pinnotheridae. Crustaceorum Catalogus, 3:
1–160. Dr. W. Junk, Den Haag.
Semper, K., 1880. Natürlichen existenzbedingungen de Thiere.
Erster theil. Leipzig: F. A. Brockhaus. Pp. 98–100.
Shen, C.-J., 1932. The Brachyuran Crustacea of North China.
Zoologica Sinica, (A)9(1): 1–320, pls. 1–10, 1 map.
Stimpson, W., 1858. Prodromus descriptionis animalium
evertebratorum quoe in Expeditione ad Oceanum Pacificum
eptentrionalem a Republica Federata Missa, Cadwaladaro
Ringgold et Johann Rodgers Ducibus, observavit et descripsit– Part V, Crustacea Ocypodoidea. Proceedings of the Academyof Natural Science, Philadelphia, 9: 93–110. (in Latin)

225

Ahyong & Ng: Pinnotherid types of Bürger (1895)
Takeda, M. & K. Konishi, 1988. Redescription of Pinnotheres
alcocki Rathbun, 1909, a commensal pea crab new to Japan.
Bulletin of the National Science Museum, Series A (Zoology),
14(3): 137–144.

226

Tesch, J. J., 1918. Decapoda Brachyura II. Goneplacidae andPinnotheridae. Siboga Expeditie Monographite, 39c1: 149–295,
pls. 7–18.
White, A., 1846. Notes on four new genera of Crustacea. Annals
and Magazine of Natural History, 18, 176–178.