Ocypode Weber, 1795

Shih, Hsi-Te, Ng, Peter K. L., Davie, Peter J. F., Schubart, Christoph D., Türkay, Michael, Naderloo, Reza, Jones, Diana & Liu, Min-Yun, 2016, Systematics of the family Ocypodidae Rafinesque, 1815 (Crustacea: Brachyura), based on phylogenetic relationships, with a reorganization of subfamily rankings and a review of the taxonomic status of Uca Leach, 1814, sensu lato and its subgenera, Raffles Bulletin of Zoology 64, pp. 139-175 : 145-147

Ocypode Weber, 1795

( Fig. 5 )

Ocypode Weber, 1795: 92 . Type species: Cancer ceratophthalmus Pallas, 1772 , by subsequent designation, see Latreille (1810: 95, 422). Gender feminine.

Ocypode Fabricius, 1798: 312 , 347. Type species: Cancer ceratophthalmus Pallas, 1772 , by subsequent designation, see Latreille (1810). Junior objective homonym of Ocypode Weber, 1795 . Gender feminine.

Ocypoda Lamarck, 1801: 149 (incorrect subsequent spelling for Ocypode Weber, 1795 ).

Monolepis Say, 1817: 155 . Type species: Monolepis inermis Say, 1817 (= Cancer quadratus Fabricius, 1787 ) by subsequent designation, see Fowler (1912). Gender feminine.

Ceratophthalma MacLeay, 1838: 64 . Type species Cancer cursor Linnaeus, 1758 , by monotypy.

Parocypoda Neumann, 1878: 26 . Type species Cancer ceratophthalmus Pallas, 1772 , by monotypy. Junior objective synonym of Ocypode Weber, 1795 . Gender feminine.

Hoplocypode Sakai & Türkay, 2013: 675 . Type species Ocypoda occidentalis Stimpson, 1860 , by original designation. Gender feminine.

Diagnosis. Body deep; carapace subquadrangular, regions poorly defined; front wide, prominently deflexed; eyestalks large, cornea occupying most of ventral surface of stalk, tip cornea sometimes with stylus, horn or prominent setal brush; antenna relatively long; inner antennal septum broad; chelipeds unequal in both sexes, palm of larger chela usually with stridulating ridge of tubercles and/or striae; first to fourth ambulatory legs relatively short, stout, dactylus fluted; pleonal locking mechanism absent; G1 typically simple distally, but may be complex in shape distally, may appear hoof-shaped in mesial view. Atlantic and Indo-Pacific.

Species included:

1. Ocypode africana De Man, 1881

= Ocypoda hexagonura Hilgendorf, 1882

= Ocypoda edwardsi Osorio, 1890

2. Ocypode brevicornis H. Milne Edwards, 1837

3. Ocypode ceratophthalmus ( Pallas, 1772)

= Cancer arenarius Toreen , in Osbeck, 1765

= Cancer caninus Herbst, 1782

= Ocipode urvillei Guérin, 1829

= Ocypoda brevicornis var. longicornuta Dana, 1852 = Ocypoda macleayana Hess, 1865

= Cancer francisci Curtiss, 1938

4. Ocypode convexa Quoy & Gaimard, 1824

5. Ocypode cordimana Latreille, 1818

= Cancer roberti Curtiss, 1938

6. Ocypode cursor ( Linnaeus, 1758)

= Cancer eques Aubert de la Chesnaye des Bois, 1759

= Ocypoda ippeus Olivier, 1804

7. Ocypode fabricii H. Milne Edwards, 1837

8. Ocypode gaudichaudii H. Milne Edwards & Lucas, 1843 9. Ocypode jousseaumei ( Nobili, 1905)

10. Ocypode kuhlii De Haan, 1835

11. Ocypode macrocera H. Milne Edwards, 1852

= Ocypode portonovoensis Prem Kumar & Tiwari, 1964

12. Ocypode madagascariensis Crosnier, 1965 13. Ocypode mortoni George, 1982

14. Ocypode nobilii De Man, 1902

15. Ocypode occidentalis Stimpson, 1860

16. Ocypode pallidula Hombron & Jacquinot, 1846

= Ocypoda laevis Dana, 1852

17. Ocypode pauliani Crosnier, 1965

18. Ocypode platytarsis H. Milne Edwards, 1852 19. Ocypode pygoides Ortmann, 1894

20. Ocypode quadrata ( Fabricius, 1787)

= Cancer arenarius Catesby, 1771 (not available name)

=? Ocypode rhombea Fabricius, 1798

= Ocypode albicans Bosc, 1801

= Monolepis inermis Say, 1817

21. Ocypode rotundata Miers, 1882

= Ocypode rotundata var. arabica Nobili, 1906 22. Ocypode ryderi Kingsley, 1880

23. Ocypode saratan ( Forskål, 1775)

= Ocypode aegyptica Gerstaecker, 1856 24. Ocypode sinensis Dai, Song & Yang, 1985 25. Ocypode stimpsoni Ortmann, 1897

Two species are nomina dubia: Ocypode laevis Fabricius, 1798 ; and Ocypode minuta Fabricius, 1798 (see Ng et al., 2008).

Remarks. The ghost crabs have been revised in detail by Sakai & Türkay (2013) with the recognition of 21 valid species, although Ng et al. (2008) listed 26 species. Hoplocypode Sakai & Türkay, 2013 , was recently established for O. occidentalis (central east Pacific coast of the Americas) based on the morphology of G1 ( Sakai & Türkay, 2013); but this taxon is not supported in our study ( Fig. 2 ). Interestingly, our tree does support two subclades of ghost crabs: O. gaudichaudii , O. occidentalis , and O. quadrata belong to a highly-supported American subclade; while all remaining species form the other, although with a lower ML support value. The three members of the American subclade share similar first gonopodal characters and may prove to be a distinct genus. However, even if they need to be placed in their own genus, Hoplocypode Sakai & Türkay, 2013 , will still be a junior subjective synonym of Monolepis Say, 1817 , whose type species is Monolepis inermis Say, 1817 (= Cancer quadratus Fabricius, 1787 ). Sakai & Türkay (2013) synonymised a number of species, notably O. sinensis Dai, Song & Yang, 1985 , with O. cordimana Latreille, 1818 . However, Huang et al. (1998) already showed they differ in a number of adult morphological characters (although juveniles are very close and hard to separate) and there are also genetic differences ( Fig. 1 ; Wong et al., 2012: fig. 9). Further evidence showing morphological and molecular differences between these two species is being prepared (Shih et al., in prep.). Further studies with more species of Ocypode than included herein will be necessary to clarify the relationships within this genus.

Figures

Fig. 5. Photographs of some species of the genus Ocypode. A, O. ceratophthalmus (Taiwan); B, O. cordimana (Dongsha, Taiwan); C, O. fabricii (Northern Territory, Australia); D, O. kuhlii (Christmas Island); E, O. occidentalis (Panama); F, O. rotundata (Iran); G. O. sinensis (Dongsha, Taiwan); H, O. stimpsoni (Taiwan).

Fig. 2. A Bayesian inference (BI) tree of the family Ocypodidae and outgroups, based on the combined 28S, 16S and COI markers. The solid circle at the node means it is strongly supported by both BI (≥ 90) and ML (≥ 70); and the open circle means only one method is strongly supported. Name of species or genus quoted means it is suggested as a synonym in this study.

Fig. 1. A Bayesian inference (BI) tree of the family Ocypodidae and outgroups, based on the combined 28S rRNA, 16S rRNA and COI markers, showing the three clades (three subfamilies), indicated in different colours. The subfamily Gelasiminae is composed of two groups, one is for all Indo-West Pacific fiddler crabs, the other for the American broad-fronted fiddler crabs.