RAFFLES BULLETIN OF ZOOLOGY 2019
Taxonomy & Systematics
RAFFLES BULLETIN OF ZOOLOGY 67: 337–351

Date of publication: 30 May 2019
DOI: 10.26107/RBZ-2019-0025
http://zoobank.org/urn:lsid:zoobank.org:pub:F42718C4-F0F4-4D75-80C3-2D155B5B8448

Two new genera of pinnotherid crabs (Crustacea: Brachyura:
Pinnotheroidea) from the Americas and the Western PacificPeter K. L. Ng1*, Shane T. Ahyong2 & Ernesto Campos3
Abstract. Orthotheres Sakai, 1969, was recently restricted to three species from the western Pacific, noting that
other species assigned to this genus from the Americas, Philippines and Palau belonged to other genera. Studiesof the non-Orthotheres material confirm that the species from the Americas should be placed in a new genus,
Mesotheres, characterised by the structure of the carapace, eyes, third maxilliped and ambulatory legs. Mesotheres
includes Pinnotheres barbatus Desbonne, in Desbonne & Schramm, 1867, Pinnotheres serrei Rathbun, 1909,
Pinnotheres strombi Rathbun, 1905, and Fabia unguifalcula Glassell, 1936. Three species from the western Pacific,
Pinnotheres glaber Bürger, 1895, Pinnotheres laevis Bürger, 1895, and Pinnotheres longipes Bürger, 1895, arereferred to a new genus, Tacitotheres, and can be distinguished by carapace form, as well as structure of the third
maxilliped and ambulatory legs.
Key words. systematics, new genera, Pinnotheridae, symbionts of gastropods, revision, Neotropics, western Pacific

INTRODUCTION

Campos’ (1989) review of the taxonomy of Orthotheresreferred Pinnotheres barbatus Desbonne, in Desbonne &Schramm, 1867, and Fabia unguifalcula Glassell, 1936,
from Mexico, to the genus, but argued that O. rathbunae
Schmitt, McCain & Davidson, 1973, O. laevis and O.
longipes should be excluded. Later, Geiger & Martin (1999)
described Orthotheres haliotidis from Australia. Ahyong& Ng (2007a) re-examined the material of Bürger (1895),
showed that O. rathbunae was referrable to Nepinnotheres
Manning, 1993a, and transferred Pinnotheres glaber Bürger,
1895 (Philippines) (treated as a subjective senior synonymof Pinnotheres impressus Bürger, 1895) and Pinnotheres
longipes Bürger, 1895 (from Philippines) back to Orthotheres.

Sakai (1969) established Orthotheres for a new species of
pinnotherid crab, O. turboe Sakai, 1969, collected from theturban shell Turbo argyrostomus Linnaeus, 1758 (Turbinidae)
from Japan and Palau. He also transferred the following
species to Orthotheres: Pinnotheres barbatus Desbonne,
in Desbonne & Schramm, 1867 (West Indies); Pinnotheres
laevis Bürger, 1895 (Philippines); Pinnotheres longipes
Bürger, 1895 (Philippines); Pinnotheres serrei Rathbun,
1909 (Puerto Rico); and Pinnotheres strombi Rathbun, 1905
(Florida). Sakai (1969: 275) listed the species as “Pinnotheres
barbatus Rathbun — Host: Turbo” but his citation was
inaccurate – he was actually referring to Rathbun’s (1918:90) record of Pinnotheres barbatus Desbonne, in Desbonne& Schramm, 1867, from Turbo. Schmitt et al. (1973)
proposed a replacement name, Orthotheres rathbunae, for
Pinnotheres barbatus Bürger, 1895, as it was preoccupied by
Pinnotheres barbatus Desbonne, in Desbonne & Schramm,
1867; probably assuming Bürger’s (1895) species was alsoan Orthotheres, like Desbonne’s (1867) taxon.

Ng & Ho (2016) restricted Orthotheres to three species
from the western Pacific associated with turban shells(Turbinidae) and abalones (Haliotidae): O. baoyu Ng &Ho, 2016, O. haliotidis and O. turboe. These three species
share a transversely ovate and wider than long carapace
with a distinct front that is clearly visible in dorsal view; anischiomerus of the third maxilliped which is subrectangular
to subovate (mesial margin gently concave); the propodus of
the third maxilliped being subcylindrical and elongate withits dorso-distal margin extending beyond the articulation
with the dactylus, which is inserted subterminally; and thedactyli of the ambulatory legs being strongly curved, almost
at right angles, with those of the second and third legs
having the ventral surface prominently concave (especiallyin females). Using these characters, they commented thatthe other species placed in Orthotheres should be referred
to two other unnamed groups.

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

Marine Invertebrates, Australian Museum Research Institute, Australian Museum,
1 William Street, Sydney, NSW 2010, Australia; and School of Biological, Earth &Environmental Sciences, University of New South Wales, Kensington, NSW 2052,
Australia; Email: shane.ahyong@austmus.gov.au
2

Facultad de Ciencias, Universidad Autónoma de Baja California, Ensenada, Baja
California, México; Email: ecampos@uabc.edu.mx
3

The first group, with all American species (O. barbatus,
O. serrei, O. strombi and O. unguifalcula), is united by afemale carapace that is also distinctly wider than long, but© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)337

Ng et al.: On two new genera of pinnotherid crabs
SYSTEMATICS

with the front strongly deflexed such that it is not visible
in dorsal view, the ischiomerus of the third maxilliped isvery elongated (mesial margin very concave), and the third
maxilliped propodus is short and subovate or subconical(not elongate) with the dactylus inserted terminally; andthe dactyli of the ambulatory legs are gently curved withthose on the second and third legs having a convex surface(not strongly hooked with the ventral surface concave) (cf.
Rathbun, 1905: unnumbered second figure; Rathbun, 1909:69, unnumbered figure; Rathbun, 1918: figs. 41, 44a, 45, pl.
19 figs. 1–4, 8–11, pl. 20 figs. 1, 2; Campos, 1989: figs. 1b,2, 3a). The second group contains the western Pacific species(O. glaber [= O. impressus], O. laevis and O. longipes),and all have a clearly rounded carapace with the frontal
margin still visible from dorsal view, the third maxilliped
dactylus is inserted subterminally or terminally, and theambulatory dactyli are asymmetrical, gently curved withthe ventral surfaces of the second and third legs rounded(not prominently hooked with the ventral surfaces concave)(cf. Bürger, 1895: pl. 9 figs. 23–25, 31; pl. 10 figs. 21–24;
Ahyong & Ng, 2007a: figs. 3E, F, 24).

Family Pinnotheridae De Haan, 1833
Mesotheres, new genus
Type species. Pinnotheres strombi Rathbun, 1905, bypresent designation.
Diagnosis. Female carapace transversely ovate, distinctly
wider than long, frontal margin strongly deflexed, not visible
in dorsal view; male carapace prominently rounded, enlarged,
slightly wider than long, frontal margin just visible fromdorsal view. MXP3 with ischium and merus completely
fused, very elongate, slender, carpus of palp ovate to round,larger than propodus and dactylus, articulating on subventral
margin, propodus short, rounded, dactylus very short, knoblike, inserted terminally on propodus; exopod very short,concealed by ischiomerus in-situ, without flagellum. P2–P5
short, bilaterally symmetrical, merus very short, P4 and P5
propodus broad, subrectangular, P5 shortest, dactylus short,
gently curved, P3 and P4 dactylus with ventral surface
convex, not excavated. Male pleon with 6 free somites andtriangular telson; G1 straight; G2 exopod absent.

The authors have since examined specimens in the lasttwo groups and confirm that the characters that have been
discussed so far are constant and valid at the generic level.
The present paper serves to formally describe two new
genera, Mesotheres, new genus, and Tacitotheres, new genus.

Constituent species. Pinnotheres barbatus Desbonne, in
Desbonne & Schramm, 1867, Pinnotheres serrei Rathbun,
1909, Pinnotheres strombi Rathbun, 1905, and Fabia
unguifalcula Glassell, 1936.

MATERIAL AND METHODS

Etymology. The name is derived from the term MesoAmerica for Central America, where most of the constituent
species occur, together with a typical pinnotherid genericsuffix “-theres”. The gender of the genus is masculine.

The terminology used in herein follows Manning (1993a) andAhyong & Ng (2007a). The following abbreviations are used:
MXP3 = third maxilliped; G1 = male first gonopod; G2 =male second gonopod; P2–P5 = pereiopods 2–5 (ambulatory
legs 1–4), respectively; s. lato = sensu lato (i.e., in the
broad sense); s. str. = sensu stricto (i.e., in the strict sense).
Specimens examined are deposited in the National Museumof Natural History, Smithsonian Institution, Washington
D.C. (USNM); and Senckenberg Museum Forschunginstitut,Frankfurt, Germany (SMF); Crustacean Collection of the
Department of Biology, University of Louisiana, Lafayette,
U.S.A (ULLZ); and the Zoological Reference Collectionof the Lee Kong Chian Natural History Museum, National
University of Singapore (ZRC).

Remarks. The four American species previously referredto Orthotheres and herein transferred to Mesotheres havea unique suite of carapace, P2–P5 and MXP3 characters.
Most characteristic is the unusual structure of the MXP3,
which has the ischiomerus prominently elongate, with the
carpus enlarged and larger than the propodus and dactylus,
and the very short exopod, which lacks a flagellum andis completely concealed by the ischiomerus (Figs. 5A, B,
6A–C, I, J, 8). In Orthotheres s. str. and Tacitotheres, newgenus, the MXP3 ischiomerus is proportionately wider, thecarpus is large but still proportionately smaller, and mostsignificantly, the exopod is large and elongate, with a distinct
flagellum, and in-situ, is not completely hidden behind theischiomerus (cf. Ng & Ho, 2016: figs. 2B, G, 5B, D, 8D).
Mesotheres also differs from Orthotheres and Tacitotheresin having a straight (versus arcuate G1) and lacks the G2
exopod (versus present in Orthotheres and Tacitotheres). In
addition, Mesotheres differs from Orthotheres in having theventral surface of the P3 and P4 dactylus gently convex as inmost pinnotherids rather than distuinctly excavated (cf. Ng& Ho, 2016: figs. 3E, G, 6J, K, 9E), and also differs from
Tacitotheres in the non-clavate female P4 and P5 propodi.

Comparative material. In addition to the material of thethree species of Orthotheres s. str. from the Indo-West Pacificexamined by Ng & Ho (2016), the following specimen
was also examined: Orthotheres turboe Sakai, 1969: ZRC
2003.0363, 1 female (12.2 × 8.9 mm), Balicasag Island,
Panglao Island, Visayas, central Philippines, host not known,coll. P.K.L. Ng, from local fishermen, 25–30 July 2003.

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Fig. 1. Mesotheres strombi (Rathbun, 1905) new combination,
male (4.5 × 4.0 mm) (USNM 172221), Florida. A, overall dorsal
habitus; B, ventral view of cephalothorax.

Fig. 2. Mesotheres strombi (Rathbun, 1905) new combination,
female (7.3 × 5.3 mm) (USNM 172221), Florida. A, overall dorsal
habitus; B, ventral view of cephalothorax.

The MXP3 of the American Bonita Campos, 2009 (typespecies Bonita mexicana Campos, 2009), Enigmatheres
Campos, 2009 (type species Fabia canfieldi Rathbun, 1918),
and Austinotheres Campos, 2002 (type species Pinnotheres
angelicus Lockington, 1877), are also similar to that ofMesotheres, new genus, but in these genera, the exopod
is elongate, visible in outer view and possess a distinct
flagellum (Campos, 2002: fig. 1D, F; Campos, 2009: figs.
1A, B, 3B, D, E).

Material examined. USNM 172221, 1 male (4.5 × 4.0 mm),
1 female (7.3 × 5.3 mm), Cedar Keys, coll. R. Merz & A.
B. Williams, 1979; USNM 122770, 1 male (12.5 × 10.2mm), 1 female (16.5 × 13.2 mm), from mantle of Florida
Horse Conch Pleuroploca gigantea (at present Triplofusus
giganteus [Kiener, 1840]) (Fasciolariidae), Cape Florida flats,
Florida, U.S.A, coll. J. Roth, 26 May 1965; ULLZ 2595, 1
female, from within spire of large Pleuroploca, St. Joseph’s
Bay, Florida, U.S.A., coll. S. & G. Graves, 25 July 1982.

We have included O. serrei (Rathbun, 1909) (from Puerto
Rico) and O. unguifalcula (Glassell, 1936) (from Mexico)
in Mesotheres as their characters fit the current diagnosisof the genus. Mesotheres serrei, was well described andfigured (Rathbun, 1909: 69, unnumbered fig.; Rathbun,
1918: 84, text-fig. 41, pl. 19 figs. 1–7) and there is no needto elaborate on its taxonomy here. Similarly, the taxonomyof M. unguifalcula, from Mexico, was treated at length by
Campos (1989: 1125, figs. 2, 3) and need not be revisited.
Campos (1989) observes that M. unguifalcula is close to M.
serrei and M. strombi.

Diagnosis. Carapace glabrous, convex in all directions.
Short pubescence on carapace frontal margin, lower surfaceof carapace, pleon, MXP3, and margins of P2–P4 except
dactyli. Eyes partially visible in dorsal view; front distinctly
deflexed, defined by convex median projection in frontal
view. P2–P4 similar, slender; P5 remarkably short, narrower
than other legs.
Remarks. The female specimens of A. strombi studied by
Rathbun (1905, 1918) had carapace width to length ratiosof 1.45, but the two females examined are only 1.28 and
1.38, respectively, possibly due to individual variation andthe state of preservation. Hernández et al. (2017) recently
noted that his Venezuelan material had a mean carapace
width to length ratio of 1.5.

Mesotheres strombi (Rathbun, 1905) new combination(Figs. 1–6)
Pinnotheres strombi Rathbun, 1905: 371, figs. 1, 2; Rathbun,
1918: 90, pl. 20 figs. 1, 2; Wass, 1955: 158; Menzel, 1956:
46; Reed, 1992: 229.
Orthotheres strombi — Sakai, 1969: 275; Schmitt et al., 1973:
27; Abele & Kim, 1986: 64, fig. 705; Ng et al., 2008: 250;
Felder et al., 2009: 1086; Hernández et al., 2017: 17, figs, 1, 2.

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Fig. 3. Mesotheres strombi (Rathbun, 1905) new combination, male (12.5 × 10.2 mm) (USNM 122770), Florida. A, overall dorsal habitus;B, frontal view of cephalothorax; C, ventral view of cephalothorax.

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Fig. 4. Mesotheres strombi (Rathbun, 1905) new combination, female (16.5 × 13.2 mm) (USNM 122770), Florida. A, overall dorsal
habitus; B, frontal view of cephalothorax; C, ventral view of cephalothorax.

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Fig. 5. Mesotheres strombi (Rathbun, 1905) new combination, male (4.5 × 4.0 mm) (USNM 172221), Florida. A, outer view of right
MXP3; B, inner view of right MXP3; C, pleon; D, anterior thoracic sternum, buccal cavity and right third MXP3; E, outer view of rightchela; F, right P2; G, right P3; H, right P4; I, right P5; J, K, left G1; L, left G2. Scales: A, B, J–L = 0.5 mm; C–I = 1.0 mm.

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Fig. 6. Mesotheres strombi (Rathbun, 1905) new combination, A–H, female (7.3 × 5.3 mm) (USNM 172221), Florida; I–P, male (12.5 ×
10.2 mm) (USNM 122770), Florida. A, I, outer view of right MXP3; C, K, outer view of carpus, propodus and dactylus of right MXP3;
B, J, inner view of right MXP3; D, outer view of right chela; E, right P2; F, right P3; G, right P4; H, right P5; L, pleon; M–O, left G1;
P, left G2. Scales: A, B, D–H, I, J, M–P = 1.0 mm; C, K = 0.5 mm; L = 2.0 mm.

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Fig. 7. Mesotheres barbatus (Desbonne, in Desbonne & Schramm, 1867) new combination, female (9.7 × 7.7 mm) (USNM 102159),
Venezuela. A, overall dorsal habitus; B, frontal view of cephalothorax; C, ventral view of cephalothorax.

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Mesotheres barbatus (Desbonne, in Desbonne &Schramm, 1867) new combination(Figs. 7, 8)
Pinnotheres barbata Desbonne, in Desbonne & Schramm, 1867: 44.
Pinnotheres barbatus — Rathbun, 1918: 88, text-fig. 44, pl. 19, figs.
8–11; Rathbun, 1933: 82; Schmitt et al., 1973: 39; Manning &Holthuis, 1981: 183; Gore, 1986: 147, table 1.
Orthotheres barbatus — Sakai, 1969: 244; Campos, 1989: 1124;
Ng et al., 2008: 250; Palacios-Theil et al., 2009: 449; Palacios
Theil et al., 2016: 5.

Material examined. USNM 102159, 1 female (9.7 × 7.7mm), zoeas and megalopa, Los Roques Islands, Venezuela,
Caribbean Sea, coll. J. Bolaños, 18 December 2000.
Diagnosis. Carapace glabrous, convex in all directions.
Margin of carapace front, under surface of carapace, pleon,
margins of chelipeds and P2–P5 strongly hirsute (except
dactyli). Eyes and front not visible in dorsal view; front
distinctly deflexed, defined by convex median projection
in frontal view. P1–P5 similar, broad, compressed; P5 notprominently shorter, narrower than other legs.

Fig. 8. Mesotheres barbatus (Desbonne, in Desbonne & Schramm,1867) new combination, female (9.7 × 7.7 mm) (USNM 102159),
Venezuela. A, outer view of right MXP3; B, inner view of right
MXP3. Scales = 0.5 mm.

Remarks. The larval development of A. barbatus, describedby Bolanõs et al. (2005), is abbreviated, with only two zoeal
stages (usually three or more in other pinnotherines) preceding
the megalopa (see also Galindo et al., 2003).

Tacitotheres, new genus

Mesotheres unguifalcula (Glassell, 1936) new
combination

Type species. Pinnotheres glaber Bürger, 1895, by original
designation.

Pinnotheres unguifalcula Glassell, 1936: 298, pl. 21 fig. 2; Schmitt
et al., 1973: 26.
Orthotheres unguifalcula — Campos, 1989: 1125, figs. 2, 3; Nget al., 2008: 250; Campos, 2018: 386, fig. 1D.

Diagnosis. Female carapace rounded, slightly wider thanlong, frontal margin visible from dorsal view; male carapace
rounded, slightly wider than long, with distinct front. MXP3
with ischium and merus completely fused, elongate; carpus
subovate, small, articulating on subventral margin; propodus
elongate, cylindrical, larger than carpus or dactylus; dactylus
slender, relatively short, inserted subterminally or terminallyon outer surface of propodus; exopod slender, long, adjacentto ischiomerus in situ, with short flagellum. P2–P5 short,bilaterally asymmetrical from left to right; merus relatively
elongate; P4 and P5 propodus clavate with distal part wider
than proximal part; P5 shortest, dactylus short, gently curved;
P3 and P4 dactylus with ventral surface convex in crosssection, not excavated. Male pleon with 6 free somites andtelson; G1 arcuate; G2 exopod distinctly longer than endopod.

Material examined. USNM, 2 females, host unknown,
Punta (Puerto) Peñasco, Sonora, México, coll. S.A. Glassell,no date; USNM 1447261, 2 females, CES IRN 11738697,
Gulf of California, Mexico, Sonora, Punta (Puerto) Penasco,
31°19′N 113°32′W, coll. S. Glassell, 2 April 1937; USNM
1447260, 1 female, CES IRN 11738696, Gulf of California,
Mexico, Sonora, Punta (Puerto) Penasco, 31°19′N 113°32′W,
coll. S. Glassell, 3 May 1935; USNM 1447262, 1 ovigerous
female, CES IRN 11738699, Mexico, Guerrero, Acapulco,
16°50′47″N 99°52′39″W, coll. E. Huffman, 22 December
1937.

Constituent species. Pinnotheres glaber Bürger, 1895,
Pinnotheres laevis Bürger, 1895, and Pinnotheres longipes
Bürger, 1895.

Diagnosis. Carapace glabrous, convex in all directions.
Carapace frontal margin glabrous, short pubescence on upper
margins of meri of P2 and P3, and anterior lower marginof carpus and propodus of P2. Eyes and front not visible
in dorsal view; front distinctly deflexed, defined by convex
median projection in frontal view. P2–P4 similar, slender;
P5 remarkably shorter, narrower than other legs.

Etymology. The name is derived from the Latin, “tacitus”, for
hidden and secret, in combination with a typical pinnotherid
generic suffix “-theres”, alluding to the fact that the three
species have not been collected for over 100 years! The
gender of the genus is masculine.

Remarks. Campos (1989) treated the taxonomy of thisspecies at length and is therefore not elaborated on here.

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Remarks. This new genus is established for three species
formerly placed in Orthotheres: Pinnotheres glaber Bürger,
1895 (type species), P. laevis Bürger, 1895, and P. longipes
Bürger, 1895. Although P. laevis and P. longipes are
currently known only from their type description (Bürger,1895), they are included in Tacitotheres, new genus, basedon the form of their MXP3 and the assumption that thefemale P4–5 propodus is clavate (these appearing, at most,weakly clavate in Burger [1895: pl. 9 figs. 25, 31]). The
P4 and P5 propodi of T. glaber are clavate (Fig. 10A, D)
but were not clearly figured as such by Bürger (1895: pl.
9 fig. 23) and we suspect a similar situation pertains to thetype figures of P. laevis and P. longipes (see Burger 1895:
pl. 9 figs. 25, 31).

The MXP3 of Tacitotheres also resembles that of Abyssotheres
Manning & Galil, 2000 (see also Komatsu & Ohtsuka, 2009)
but differs in possessing a larger palp with the propodus
larger than the carpus (Fig. 10E, H, M), instead of a short
palp with the carpus larger that propodus as in Abyssotheres(cf. Manning & Galil, 2000: fig. 1a; Komatsu & Ohtsuka,
2009: fig. 3F). Tacitotheres further differs in the eyes ofthe female that are not dorsally visible (Figs. 9A, B, 10A)
(versus clearly well visible in dorsal view in Abyssotheres),the propodi of P2–P5 are clavate (Fig. 10A) (subquadrate
in Abyssotheres) and the ambulatory legs are asymmetrical(Figs. 9, 10A) (symmetrical in Abyssotheres) (cf. Manning &Galil, 2000: 66; Komatsu & Ohtsuka, 2009: figs. 1A, 2D–G).
The American Bonita and Enigmatheres also have a clavate
P5 propodus (Campos, 2009: figs. 1E, 2C), but have a
proportionately more elongate MXP3 with an enlarged
carpus. Another American genus, Raytheres Campos, 2004
(type species Pinnotheres clavapedatus Glassell, 1935) also
has clavate ambulatory propodi, but these are on P2 and P3
instead (see Campos, 2002: fig. 5E).

Of Pinnotheres laevis Bürger, 1895, Campos (1989)
already commented that it is not a species of Orthotheres
because of its carapace proportions and relative lengths of
the articles of the third maxilliped palp, and suggested it
be referred to Pinnotheres s. lato instead. The transfer ofP. laevis and P. longipes to Tacitotheres addresses someof these concerns. Nevertheless, the correct taxonomy ofthese species is seriously compromised by the loss of typespecimens of both species. Until P. laevis and P. longipes
can be evaluated based on specimens, they are tentatively
placed in Tacitotheres.

Tacitotheres glaber (Bürger, 1895) new combination(Fig. 9, 10)
Pinnotheres glaber Bürger, 1895: 379, pl. 9: fig. 23, pl. 10:
fig. 21; Tesch, 1918: 248; Miyake, 1939: 221, 241; Silas &Alagarswami, 1967: 1199, 1216; Ng & Manning, 2003: 902.
Pinnotheres impressus Bürger, 1895: 380, pl. 9: fig. 24, pl. 10:
fig. 23; Tesch, 1918: 249, 255; Silas & Alagarswami, 1967:
1200, 1216.
Orthotheres glaber – Ahyong & Ng, 2007a: 218, fig. 24; Ng etal., 2008: 250.

The MXP3 of Tacitotheres, including the presence of a welldeveloped exopod with a short flagellum, closely resembles
that of Orthotheres s. str. (cf. Ng & Ho, 2016: figs. 2B, C, G,
5B–E, 8D–F). The ambulatory legs of Tacitotheres, however,are distinctively more elongate and slender (Fig. 10 A, F)(versus short and stout in Orthotheres s. str., cf. Ng & Ho,
2016: figs. 3A, B, D, F, H, J–M, 6A–I, K, 9A–D, F–I) andthe ventral surface of the P3 and P4 dactylus is convex incross-section (versus distinctly concave in Orthotheres s. str.,cf. Ng & Ho, 2016: figs. 3E, G, 6J, K, 9E). In addition, theG2 of T. glaber has a well-developed and very long exopod(Fig. 10L) (exopod four times endopod length versus aslong as or shorter than the endopod in Orthotheres s. str.;
Ng & Ho, 2016: fig. 5M). The female P4 and P5 propodus
is distinctly clavate in shape (i.e. distal part wider thanproximal part) in T. glaber (Fig. 10A), a feature not present
in other species that have been associated with Orthotheres
s. str. or Mesotheres, new genus, and that we have examined(except possibly O. laevis and O. longipes, see below).
Species of Orthotheres s. str. have a more symmetrical P4
and P5 propodus (cf. Ng & Ho, 2016: figs. 3A, B, L, M,
6C, D, G, H, 9–I) than Tacitotheres as does Mesotheres(Figs. 5H, I, 6G, H).

Material examined. Lectotype: USNM 32431, ovigerous
female (7.8 × 6.5 mm), Palau, from Tapes turgida, coll. C.
Semper. Paralectotypes: SMF-ZMG 173, Go304a, 6 males(3.4 × 3.1 to 4.8 × 4.3 mm), 4 females (3.5 × 3.5 to 8.3 ×
7.4 mm), same locality as lectotype.
Diagnosis. Female: Carapace subcircular, slightly wider thanlong, front faintly projecting, convex. Eyes small, not visible
in dorsal view. MXP3 with propodus more than twice aslong as high, apex rounded; dactylus inserting subdistally,outreaching propodal apex; ischiomerus exceeding twice
length, outer margin convex, inner margin with proximal¾ concave, angular at distal quarter; exopod outermargin convex, inner margin relatively straight, flagellum
2-segmented. Cheliped with dactylus about 2/3 palm length;
palm 2½ times as long as high, mesioventral margin setose;
dactylus occlusal margin finely setose, with large proximal
tooth; pollex occlusal margin sinuous, sparsely setose.
Ambulatory 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 andP5 propodus with sparse ventral setae. Pleon extending
to buccal region and covering bases of ambulatory legs.
Male: Carapace rounded, wider than long, front projecting,medially emarginate. Eyes visible in dorsal view. MXP3 as

The general carapace shape and left-right symmetry of
the ambulatory legs of Tacitotheres resembles those of
Pinnotheres Bosc, 1802, and Nepinnotheres Manning,
1993a, but the structure of the MXP3 differs significantly.
In Tacitotheres, the MXP3 dactylus is short and inserted
clearly subterminally on the outer surface of the propodus,
with the propodus the longest segment, and the carpus shortand not enlarged (Fig. 10E, H, M).

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Fig. 9. Tacitotheres glaber (Bürger, 1895) new combination, A, paralectotype male (4.6 × 4.2 mm) (SMF-ZMG 173); B, C, lectotype
ovigerous female (7.8 × 6.5 mm) (USNM 32431), Palau. A, B, overall dorsal habitus; C, ventral surface of cephalothorax.

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Ng et al.: On two new genera of pinnotherid crabs

Fig. 10. Tacitotheres glaber (Bürger, 1895) new combination, A–E, lectotype female (7.8 × 6.5 mm) (USNM 32431), Palau; F–I,
paralectotype male (4.0 × 3.6 mm) (SMF-ZMG 173); J–M, paralectotype male (4.6 × 4.2 mm) (SMF-ZMG 173). A, F, dorsal overall
habitus; B, sternum, anterior margin; C, anterior cephalothorax; E, H, right MXP3; D, G, right chela, outer view; I, J, male pleon; K,
left G1, pleonal view; L, left G2, pleonal view, with long exopod; M, left MXP3. A, D–I, modified after Ahyong & Ng (2007a: fig. 24).
Scale: A = 2 mm; B–D, F–G, I, J = 1.0 mm; E = 0.5; H, K, L, M = 0.25 mm.

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dactyl-propodal articulation. Like Ostracotheres, which wasrecently shown to comprise several disparate genera despite
the similar MXP3 palp segmentation (Ahyong, 2018), theOrthotheres-like genera may prove to be dispersed throughout
the pinnotherine phylogeny.

in female, but sometimes 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. Ambulatory
legs dorsally and ventrally unarmed, length asymmetricalfrom left to right; relative lengths P3>P2>P4>P5. Dactyli
shorter than half-length of respective propodi; P2–P4 dactyli
subequal in length and slightly longer than P5 dactylus. P3
and P4 propodus with dorsal and ventral setae; carpus withdiagonal setal row. Pleon widest at somites 3 and 4, tapering
distally to evenly rounded telson. G1 slender, arcuate. G2
with exopod distinctly longer then endopod. (Modified from
Ahyong & Ng, 2007a: 218).

Campos (2009: 42) highlighted the presence of a protuberance
on the mesioventral margin of the “basal antennal article”
as a useful phylogenetic character within Pinnotheridae
and which may be a key diagnostic feature for the
Pinnotherinae. The protuberance is evident in Orthotheres andMesotheres, but indistinct in Tacitotheres. Unlike Orthotheres
and Mesotheres, which are associated with gastropods,
Tacitotheres is associated with bivalves. However, amonga wider suite of genera that we have assessed for the formof the antennal articles, including Alain Manning, 1998 (seealso Ahyong & Ng, 2008), Buergeres Ng & Manning, 2003,
Pinnotheres Bosc, 1802, Nepinnotheres Manning, 1993a,
Arcotheres Manning, 1993a, Holotheres Ng & Manning,
2003, Viridotheres Manning, 1996, Serenotheres Ahyong &Ng, 2005, and Visayeres Ahyong & Ng, 2007b, no patternin relation to somatic morphology or host is yet evident.
We note that the “basal antennal article” referred to by
Campos (2002, 2009) represents antennal article 1 and is
not equivalent to the “basal antennal article” frequently usedin brachyuran nomenclature, especially in the taxonomy of
Majoidea (Davie et al., 2015: 64). The “basal antennal article”
in more common usage among brachyurans, rather thanreferring to article 1, refers to the quadrate or subquadrate
structure just below the orbit, which is composed of the fusedarticles 2 and 3. Antennal article 1 is the osmoregulatory
article, which contains the green gland and its opening, thenephridiopore. In many crabs, article 1 is often small oreven indistinguishably fused to the surface of the epistome,
but the presence of the nephridiopore indicates its position.
In all the pinnotherids examined (see also Campos, 2002,
2009; Ahyong & Ng, 2005, 2007a, b, 2008; Ng & Meyer,2016), the fused articles 2 and 3 are usually demarcated byshallow grooves and article 1 is also immovably fused but
positioned very low on the epistome such that its lower
margin intrudes into, and forms part of, the margin of the
buccal cavity. This lower margin of article 1 varies fromindistinct to bluntly angular, appearing as a short lobe or
protuberance, being most pronounced mesially. Among thespecies examined, the condition of this protuberance showsno obvious taxonomic pattern within or between genera.
In Orthotheres s. str., the mesioventral angle of antennal
article 1 is slightly projected and tubercle-like, with asmall pore at the tip, almost certainly the nephridiopore.
This mesioventral position of the presumed nephridiopore,readily observed in larger specimens of Nepinnotheres andArcotheres, this may well be the normal position in most,
if not all, pinnotherines (and pinnotherids). Using primarily
molecular data, Palacios-Theil et al. (2016) recognised a
Pinnotherinae (based mostly on American representatives,
including Mesotheres barbatus), somewhat different fromthe system proposed by Campos (2009) which emphasised
morphological characters like the presence or absence of a
protuberance on the inner ventral edge of antennal article 1
as well as known larval features. The location of antennal

Remarks. It is somewhat surprising that T. glaber is known
only from the type material given that it was originallyreported from venerid clams, which are widely collected
for human consumption. The original host, Tapes turgida(Lamarck, 1818) is currently a junior synonym of T.
conspersus (Gmelin, 1791). An unusual aspect of male T.
glaber is the size of the G2 exopod and pleonal form. The
G2 exopod of T. glaber is remarkable as it is the largest
known of any pinnotherid (relatively), being about quadruple
the length of the endopod (Fig. 10L). A large G2 exopod
is also present in Viridotheres takedai Ahyong, Komai &Watanabe, 2012, and V. kupang Ahyong, 2019, but the exopod
is shorter than three times the length of the endopod. Of noteis also the form of the male pleon. Unusually, the male pleon
in T. glaber is narrowest in the smallest specimens (Fig.
10I) becoming wider with increasing body size (Fig. 10J),
potentially becoming more female-like. The significance of
this phenomenon is presently unknown and must await further
collections allowing assessment of developmental changes.
DISCUSSION
It is noteworthy that many of the species that havebeen previously placed in Orthotheres are associatedwith gastropod hosts rather than bivalves, which is themore common host type among pinnotherids. Species
in Orthotheres s. str. as well as Mesotheres, new genus,
live in vetigastropods (Haliotis, Cittarium, Turbo) andcaenogastropods (Strombus, Triplofusus), and this habit
may be a consequence of convergence rather than shared
phylogenetic history. The record of an “Orthotheres” species
in a chiton by Melzer & Schwabe (2008) from Chile wasrecently corrected by Campos (2018), who noted that it
was probably a species of Calyptraeotheres Campos, 1990,
instead.
Phylogenetic relationships among the Orthotheres-like genera
are yet to be studied and to date, only Mesotheres strombi and
M. barbatus, have been used as exemplars in phylogenetic
analyses (Pohle & Marques, 1998; Palacios-Theil et al.,2009, 2016). Most of the Orthotheres-like genera, however,appear to have little in common apart from the similar MXP3349

Ng et al.: On two new genera of pinnotherid crabs
article 1, on the margin of the buccal cavity, may prove
to be a synapomorphy of Pinnotheridae, but until morecomprehensive phylogenetic and comparative morphologicalstudies can be completed, the taxonomic and phylogenetic
value of this character remains unclear.

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ACKNOWLEDGMENTS
The authors thank Rafael Lemaitre and Karen Reed (USNM)
for their kind help in allowing access to their material; aswell as to Angelika Brandt (SMF) for expediting a loanof the type material of Orthotheres glaber. We are deeply
appreciative of the many helpful comments from Tohru
Naruse and Kooichi Konishi, as well as Jose Mendoza, in
getting this manuscript to press.
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