Ocypodidae Rafinesque, 1815

Family Ocypodidae Rafinesque, 1815 View in CoL Subfamily Gelasiminae Miers, 1886

Remarks: Traditional understanding of distinct lineages of ghost crabs (“ Ocypodinae ”) and fiddler crabs (“ Ucinae ”) has been challenged, and “fiddler crabs” (or Uca sensu lato) have been shown to be paraphyletic ( Shih et al. 2016a). Revised classification presented in Shih et al. (2016a), showing Ocypodinae Rafinesque, 1815 now containing ghost crab Ocypode Fabricius, 1798 (circumtropical), Uca Leach, 1814 sensu stricto (Americas) and Afruca Crane, 1975 (eastern Atlantic region) (the latter two being fiddler crab genera), and Gelasiminae Miers, 1886 accommodating all other fiddler crab genera, under which all subgeneric ranks raised to full generic status, is herein followed.

Austruca lactea (De Haan, 1835) #

Gelasimus lacteus View in CoL — Stimpson 1858b: 100; Stimpson 1907: 108; Shen 1940a: 231.

Gelasimus lactens — Koelbel 1897: 717.

Uca lactea View in CoL — Gordon 1931: 528; Hill et al. 1975: 65; Morton 1976: 102, fig. 55g; Hill et al. 1978: 92; Morton 1979a: 118, figs. 6.18o, 6.19; Morton and Morton 1983: 153, fig. 10.3(4); Choi 1992: 901; Jones and Morton 1994: 28, fig. 6A–I, pls. 2G, H, 3E, F; BKK Chan and Caley 2003: 92, 2 unnumb. figs.; Fong et al. 2005: 56, 1 unnumb. fig.; WPW Kwok and Tang 2006: 4, figs. 10, 11; So and Lui 2007: 31, 5 unnumb. figs.; Shih et al. 2010b: 10.

Uca (Celuca) lactea lactea View in CoL — Crane 1975: 300, figs. 19A, 54J, JJ, 69E, pl. 40A, B.

Uca (Celuca) lactea View in CoL — SY Lee and Leung 1999: 71.

Uca lactea annulipes View in CoL — Tam and Wong 2000: 117, 2 unnumb. figs.

Austruca lactea View in CoL — Bravo et al. 2021: tab. 2.

Taxonomy: Sakai (1939: as Gelasimus ), Crane (1975: as U. (Celuca) lactea lactea ), Naderloo et al. (2010: as U. ( Austruca )), Shih et al. (2015).

Type: Lectotype ò RMNH D 254, and paralectotypes in RMNH, SMF, ZMG, NHMD, MNHN (designated by Crane 1975; see also Yamaguchi and Baba 1993, Fransen et al. 1997, Naderloo et al. 2010).

Type locality: Japan, locality unspecified.

Distribution: South China Sea and East Asia: northern Vietnam, South China, Taiwan, Korea and Japan ( Shih et al. 2010b 2022).

Habitat: Intertidal; in burrows on mudflats, preferring coarser substrates.

Gelasimus borealis ( Crane, 1975) View in CoL #

Gelasimus vocans View in CoL — Stimpson 1858b: 99 [part: Hong Kong material only]; Stimpson 1907: 104 [part: Hong Kong material only]; Koelbel 1897: 717. [not Cancer vocans Linnaeus, 1758 ].

Uca marionis View in CoL — Gee 1926: 165 [list]; Morton 1979a: 118, fig. 6.17.

Uca marionis var. nitidus — Gordon 1931: 528.

Gelasimus marionis View in CoL — Shen 1940a: 232.

Gelasimus marionis nitidus View in CoL — Shen 1940a: 232.

Uca (Thalassuca) vocans borealis Crane, 1975: 90 View in CoL , fig. 64A; SY Lee and Leung 1999: 71.

Uca vocans View in CoL — Hill et al. 1975: 48; Morton 1976: 102; Hill et al. 1978: 92; Morton 1979a: 121; Melville and Morton 1983: 11, 1 unnumb. fig. (13); Morton and Morton 1983: 184, fig. 11.11(5). [not Cancer vocans Linnaeus, 1758 ].

Uca gaimardi View in CoL — Hill et al. 1975: 77; Hill et al. 1978: 107; Hill and Phillipps 1981: 152, pl. 51a. [not Gelasimus Gaimardi H. Milne Edwards, 1852 View in CoL = G. crassipes White, 1847 View in CoL ].

Uca gaimardii — Morton 1976: 103. [not Gelasimus Gaimardi H. Milne Edwards, 1852 View in CoL = G. crassipes White, 1847 View in CoL ].

Uca borealis View in CoL — Jones and Morton 1994: 22, fig. 4A–I, pls. 2E, F, 3A, B; BKK Chan and Caley 2003: 92, 3 unnumb. figs.; Fong et al. 2005: 56, 1 unnumb. fig.; WPW Kwok and Tang 2006: 4, figs. 6, 7; So and Lui 2007: 30, 4 unnumb. figs.; Shih et al. 2010a: 52; 2010b: 8.

Uca vocans vocans View in CoL — Tam and Wong 2000: 118, 2 unnumb. figs. [not Cancer vocans Linnaeus, 1758 ].

Gelasimus borealis View in CoL — Bravo et al. 2021: tab. 2.

Taxonomy: Crane (1975: as U. (Thalassuca) vocans borealis ), Shih et al. (2015).

Type: Holotype ò USNM 137669 About USNM , and paratypes in USNM.

Type locality: Hong Kong.

Distribution: South China Sea and East Asia: northern Vietnam, South China, Taiwan and Japan ( Shih et al. 2010b 2022).

Habitat: Intertidal; on mudflats, prefers coarse or gritty sand.

Remarks: So far seven species have been recognized in the “ Uca vocans species complex” ( Crane 1975; Shih et al. 2010a; Rosenberg 2019), while only two have been reliably reported from coasts of South China —the locally common G. borealis from Guangxi, Hainan to Fujian, whereas G. vocans sensu stricto ascertained from Hainan, with northern distributinal limit unclear ( Shih et al. 2010b). As such, older records of vocans , including those by Stimpson and Koelbel, might instead represent, or at least include the locally common G. borealis .

Gelasimus marionis Desmarest, 1823 View in CoL was indicated by Holthuis (1959) to be a junior synonym of Cancer vocans Linnaeus, 1758 . Shen (1940a) reported two distinct forms, namely G. marionis View in CoL and G. m. nitidus View in CoL , both citing the work by Alcock (1900b). In the dichotomous key provided by Alcock, these two forms are distinguished by form of crest on internal surface of male cheliped, and shape of pollex along cutting margin (p. 353, Alcock 1900b). This distinction, however, reflects simultaneous brachychelous and leptochelous conditions commonly found in a single population of Gelasimus vocans View in CoL sensu lato, and we place both entries under G. borealis View in CoL .

As for “ U. gaimardii ”, this name is now considered synonymous with Paraleptuca crassipes ( Crane 1975) View in CoL . However, Hill and Phillipps (1981, pl. 51a) presented a peculiar illustration under this name that seems to show features of two fiddler crab species: the form of male cheliped, externally tubercular and pollex scalloped, corresponds with G. borealis View in CoL , but the grayish carapace bearing dark transverse bands (discernable in the printed copy of their book), resembles P. splendida View in CoL . We place this record under G. borealis View in CoL herein.

Paraleptuca splendida (Stimpson, 1858) #

Gelasimus splendidus Stimpson, 1858b: 99 View in CoL ; Alcock 1900b: 355;

Stimpson 1907: pl. 14(2). Uca splendida View in CoL — Gee 1926: 165 [list]. Uca gaimardi View in CoL — Gordon 1931: 528. Gelasimus gaimardi View in CoL — Shen 1940a: 232. Uca (Amphiuca) chlorophthalmus crassipes View in CoL — Crane 1975: 101 [part:

Hong Kong material only], figs. 13G, 14B, BB; SY Lee and

Leung 1999: 71. Uca chlorophthalmus crassipes — Choi 1992: 901; Tam and Wong

2000: 117, 2 unnumb. figs. Uca chlorophthalmus — Morton and Morton 1983: 184, fig. 10.3(3). Uca chlorophthalmus — SY Lee 1993: tab. 1. Uca crassipes — Jones and Morton 1994: 26, fig. 5A–J, pl. 3C, D;

BKK Chan and Caley 2003: 92, 2 unnumb. figs.; Fong et al.

2005: 57, 1 unnumb. figs.; WPW Kwok and Tang 2006: 4, figs. 8,

9; So and Lui 2007: 32, 4 unnumb. figs.; Shih et al. 2010b: 9. Uca chloropthamalus — BKK Chan 2001: 24, fig. 1b. Uca (Paraleptuca) splendida — Shih et al. 2012: 34, figs. 2–4, 6, 7C. Paraleptuca splendida — Bravo et al. 2021: tab. 2.

Taxonomy: Shih et al. (2012: as Uca (Paraleptuca) ; 2015).

Type: Neotype ò ZRC 2012.0143 View Materials (designated by Shih et al. 2012).

Type locality: Tai Tam, Hong Kong (neotype).

Distribution: South China Sea and East Asia: Central Vietnam to South China, Taiwan, and Ryukyus ( Japan) ( Shih et al. 2012 2016b 2022).

Habitat: Supratidal to intertidal; in burrows on salt marshes, substrates of coarse sands and rock fragments, rather compact, covered by coastal vegetation.

Remarks: Crane (1975) recognized two distinct subspecies in her Uca (Amphiuca) chlorophthalmus : chlorophthalmus from the western Indian Ocean along eastern Africa, and crassipes from the western and central Pacific ( Shih et al. 2013). Gelasimus gaimardi H. Milne Edwards, 1852 is a junior synonym of G. crassipes White, 1847 according to Crane (1975). Among material identified as crassipes, Crane (1975) noted in some larger specimens from Hong Kong, the morphology of distal tip of the G1 was aberrant being “strikingly longer”, and suggested G. splendidus Stimpson, 1858 “would certainly have been kept to designate at least a subspecies” (p. 99). She, however, observed too much variation, especially in smaller specimens, to be sure. Stimpson’s (1858b) species was eventually resurrected by Shih et al. (2012) based on morphological and molecular approaches.

Tubuca acuta (Stimpson, 1858) #

Uca mani — Gordon 1931: 528. Gelasimus manii — Shen 1940a: 232. Gelasimus urvillei View in CoL — Shen 1940a: 232. Uca (Deltuca) acuta acuta View in CoL — Crane 1975: 25, fig. 61B, pl. 1E–H. Uca acuta View in CoL — SY Lee 1993: tab. 1; Jones and Morton 1994: 12, fig.

1A–K, pls. 1A, B, 2A, B; So and Lui 2007: 27, 3 unnumb. figs.;

Shih et al. 2010b: 3, fig. 2A–D. Uca (Deltuca) acuta — SY Lee and Leung 1999: 70. Tubuca acuta — Bravo et al. 2021: tab. 2.

Taxonomy: Crane (1975: as U. (Deltuca) acuta acuta ), Shih et al. (2010b: as Uca ; 2015).

Type: Neotype ò USNM 137665 About USNM (designated by Crane 1975).

Type locality: Castle Peak area, Kowloon, Hong Kong (neotype).

Distribution: South China Sea and East Asia: northern Vietnam, South China, and Taiwan ( Shih et al. 2015 2022).

Habitat: Intertidal; on estuarine mudflats, often associated with vegetation.

Remarks: Uca manii Rathbun, 1909 was a name given to replace the record of Gelasimus acutus reported from the Mergui Islands by De Man (1888b), which later synonymized under Gelasimus forcipatus Adams and White, 1849 by Crane (1975).

Gelasimus urvillei H. Milne Edwards, 1852 View in CoL is shown to be an eastern African species (Shih et al. 2018), despite type locality listed “Vanikoro” (= now Vanuatu), which probably erroneous ( Crane 1975). Both species are unlikely to be present in Hong Kong, and we follow Shih et al. (2010b) in considering these as U. acuta View in CoL (now Tubuca View in CoL ).

Tubuca arcuata (De Haan, 1835) View in CoL #

Gelasimus arcuatus View in CoL — Shen 1940a: 231.

Uca (Deltuca) arcuata View in CoL — Crane 1975: 44, figs. 8C, 9C, 61J, pl. 5A–F; SY Lee and Leung 1999: 70.

Uca arcuata View in CoL — Melville and Morton 1983: 11, 1 unnumb. fig. (13); Morton and Morton 1983: 184, fig. 11.11(6), pl. 19H; SY Lee 1993: tab. 1; Jones and Morton 1994: 13, fig. 2A–J, pls. 1C, D, 2C, D; Tam and Wong 2000: 116, 2 unnumb. figs.; Fong et al. 2005: 56, 1 unnumb. fig.; WPW Kwok and Tang 2006: 4, figs. 4, 5; So and Lui 2007: 28, 3 unnumb. figs.; Shih et al. 2010b: 7, fig. 2E, F.

Uca acuta View in CoL — WPW Kwok and Tang 2006: 4, fig. 2.

Tubuca arcuata View in CoL — Bravo et al. 2021: tab. 2.

Taxonomy: Shen (1932b), Crane (1975: as U. (Deltuca) arcuata ), Shih et al. (2015 2016b).

Type: Lectotype ò RMNH D 243, and paralectotypes in RMNH, NHM, NHMD, MNHN (designated by Crane 1975; see also Yamaguchi and Baba 1993; Fransen et al. 1997).

Type locality: Japan, locality unspecified.

Distribution: South China Sea and East Asia: northern Vietnam, coasts of China, Taiwan, Korea and Japan ( Shih et al. 2015 2016b 2022).

Habitat: Intertidal; on estuarine mudflats, substrates of soft mud.