Tubuca Bott, 1973

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 : 159

Tubuca Bott, 1973 , status nov.

( Figs. 10G, H , 11A–H , 12A )

Tubuca Bott, 1973b: 322 . Type species: Gelasimus urvillei H. Milne Edwards, 1852 , by original designation. Gender feminine.

Australuca Crane, 1975: 62 . Type species: Gelasimus bellator Adams & White, 1849 , by original designation. Gender feminine.

Deltuca Crane, 1975: 21. Type species: Gelasimus forcipatus Adams & White, 1849 , by original designation. Gender feminine.

Diagnosis. Medium- to large-sized species (carapace width about 15–35 mm in adults); dorsal carapace surface always with posterolateral striae in female, absent in males; front narrow; cornea round; eyestalks slender; adult male major cheliped very large, right- or left-handed, pollex sometimes with ventral carina, outer surface of major manus with small to large tubercles, carpus with antero-dorsal area flattened to facilitate chela flexion, setae on merus of minor cheliped short, stiff; male pleonites free; pleonal locking mechanism absent; no setae on lateral margins of posterior stem region of urocardiac ossicles in gastric. Indo-west Pacific.

Species included:

1. Tubuca acuta ( Stimpson, 1858)

2. Tubuca australiae ( Crane, 1975)

3. Tubuca arcuata (De Haan, 1835)

= Uca brevipes H. Milne Edwards, 1852

4. Tubuca bellator ( White, 1847)

5. Tubuca capricornis ( Crane, 1975)

= Uca pavo George & Jones, 1982

6. Tubuca coarctata (H. Milne Edwards, 1852)

= Uca rathbunae Pearse, 1912

= Uca ischnodactylus Nemec, 1939

=? Uca mearnsi Rathbun, 1913

=? Gelasimus thomsoni Kirk, 1881

7. Tubuca demani ( Ortmann, 1897)

= Uca zamboangana Rathbun, 1913

8. Tubuca dussumieri (H. Milne Edwards, 1852) (nomen protectum)

= Gelasimus caerulens Adams , in Belcher, 1848 (nomen oblitum)

= Gelasimus dubius Stimpson, 1858

9. Tubuca elegans ( George & Jones, 1982)

10. Tubuca flammula ( Crane, 1975)

11. Tubuca forcipata (Adams & White, 1849)

= Uca rubripes Estampador, 1937

= Uca manii Rathbun, 1909

11. Tubuca hirsutimanus ( George & Jones, 1982) 12. Tubuca longidigitum (Kingsley, 1880)

13. Tubuca paradussumieri (Bott, 1973)

= Uca (Deltuca) dussumieri spinata Crane, 1975 14. Tubuca polita ( Crane, 1975)

15. Tubuca rhizophorae ( Tweedie, 1950)

16. Tubuca rosea ( Tweedie, 1937)

17. Tubuca seismella ( Crane, 1975)

18. Tubuca signata ( Hess, 1865)

19. Tubuca typhoni ( Crane, 1975)

20. Tubuca urvillei (H. Milne Edwards, 1852)

Remarks. The molecular analyses ( Fig. 2 ) reveal one major clade containing all the Tubuca and Australuca species. Although the Australuca species do form a distinct subclade, their inclusion causes Tubuca , as presently conceived, to become paraphyletic. Thus, in order to maintain Australuca as a separate genus, we would have to further split Tubuca and this seems unjustified at this time. We therefore treat Australuca Crane, 1975 , as a junior synonym of Tubuca Bott, 1973 , as the latter has nomenclatural priority. This revised concept of Tubuca makes it the most speciose genus in the IWP, with 21 species. Although the genus is widely-distributed in the IWP, it does not extend as far as the eastern margin of Polynesia, where only Paraleptuca and Gelasimus species occur ( Poupin & Juncker, 2010; Fig. 4 ).

Figures

Fig. 10. Photographs of some species of the genera Minuca, Paraleptuca, Petruca and Tubuca. A, M. herradurensis (Panama); B, M. rapax (Brazil); C, Pa. chlorophthalmus (East Africa); D, Pa. crassipes (Donghsa, Taiwan); E, Pa. splendida (Penghu, Taiwan); F, Pe. panamensis (Costa Rica); G, T. arcuata (Hainan, China); H, T. bellator (Labuan, Malaysia). C, courtesy of S. Cannicci.

Fig. 11. Photographs of some species of the genus Tubuca. A, T. coarctata (Taiwan); B, T. dussumieri (Penghu, Taiwan); C, T. elegans (New Territory, Australia); D, T. forcipata (Sarawak, Malaysia); E, T. polita (Northern Territory, Australia); F, T. rhizophorae (Johor, Malaysia); G, T. rosea (Selangor, Malaysia); H, T. signata (New Territory, Australia). C, E, H, courtesy of P. Backwell.

Fig. 12. Photographs of some species of the genera Tubuca, Xeruca and Ucides. A, T. urvillei (Ranong, Thailand); B, X. formosensis (Taiwan); C, D, U. cordatus (Brazil).

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. 4. The distribution of 11 genera of fiddler crabs, mainly based on Crane (1975) and the website (http://www.fiddlercrab.info/index. html; Rosenberg, 2014).