Uca (Austruca) annulipes (H. Milne Edwards, 1837 )

Naderloo, Reza, Türkay, Michael & Chen, Hui-Lian, 2010, Taxonomic revision of the wide-front fiddler crabs of the Uca lactea group (Crustacea: Decapoda: Brachyura: Ocypodidae) in the Indo-West Pacific, Zootaxa 2500, pp. 1-38 : 7-11

Uca (Austruca) annulipes (H. Milne Edwards, 1837)

( Figs. 2 a–h, 3a–e, 4b, 12a, b, c)

Gelasimus annulipes H. Milne Edwards 1837: 55 .

Gelasimus lacteus — Krauss 1843: 14, 39. — Alcock 1900: 355 [specimens from Karachi] [not Ocypode (Gelasimus) lactea De Haan, 1835 ]

Gelasimus annulipes — White 1847: 36. — Dana 1852: 317. — H. Milne Edwards 1852: 149, pl. 4, fig. 15. — Heller 1865: 38. — Hilgendorf 1869: 85; 1879, 803. — Hoffmann 1874: 18. — Miers 1879: 488; 1880: 310. — Kingsley 1880: 148. — Richters 1880: 155. — De Man 1880: 69; 1887a: 118; 1887b, 353; 1891: 22–23 (in key), 39; 1895: 577; 1902: 483, pl. 8, figs. 5, 6. — Lenz & Richters 1881: 423. — Miers 1886: 244. — Alcock 1892: 415; 1900: 352 (in key), 353–355. — Henderson 1893: 388. — Ortmann 1894a: 758, 759; 1894b: 59, 67. — Alcock & Anderson 1894: 202. — Zehnter 1894: 178. — Lenz 1905: 365; 1910, 558. — Sewell 1913: 339, 344. — Bouvier 1915: 301. — Kemp 1915: 221, 222; 1918: 227. — Gordon 1934: 10. — Tweedie 1937: 141, fig. 1a. — T. Sakai 1940: 32. — Chhapgar 1957: 508 –509, pl. 13, figs. i–o.

Gelasimus annulipes var. lacteus — Ortmann 1894a: 759.

Uca annulipes — Nobili 1899a: 274; 1899b: 518; 1903: 20. — Doflein 1899: 193. — Crosnier 1965: 117, figs. 204, 206, 207, 212, 213. — Lanchester 1900a: 754; 1900b: 580; 1901: 549. — Schenkel 1902: 580. — Borradaile 1907: 69; 1910: 408. — Rathbun 1910: 322. — Roux 1917: 16. — Stebbing 1917: 16. — Symons 1920: 309. — Balss 1922: 142. — Boyce 1924: 250. — Maccagno 1928: 35. — T. Sakai 1936: 170; 1939: 616; 1940: 28. — Miyake 1936: 511; 1938: 109; 1939: 190, 222, 241, pl. 16, fig. 2. — Panikkar & Aiyar 1937: 295, 301. — Suvatti 1938: 74. — Chace 1942: 202. — Vatova 1943: 24. — Buitendijk 1947: 280. — Barnard 1950: 97, 98, figs. 18g –i, 19e. — Tweedie 1950: 356. — Sankarankutty 1961: 113. — Forest & Guinot 1961: 141. — Claude 1964: 11. — Pillay & Nair 1971a: 153; 1971b: 161. — Rajendran 1972: 378. — Naiyanetr 1980: 43 (in list). — Adiwiryono et al. 1984: 253, 256, 257 (all in list). — Macia et al. 2001: 216. — Ng & Davie 2002: 378. — Skov et al. 2002: 727, 728. — Lee & Lim 2003: 188. — Lim 2004: 1246 –1251. — Litulo 2004: 87; 2005: 283. — Lim et al. 2005: 111 –114. — Ghory & Siddiqui 2006: 162. — Naiyanetr 2007: 131 (in list). — Shih et al. 2009: 377.

Uca lactea — Pesta 1911: 22 –23 [erroneously spelled as U. lactaea ]. — Tesch 1918: 39. — Maccagno 1928: 29, fig. 15. — Barnard 1950: 96, 97. [not Ocypode (Gelasimus) lactea De Haan, 1835 ].

Uca lacteus — Stebbing 1917: 16, pl. 4. [not Ocypode (Gelasimus) lactea De Haan, 1835 ].

Austruca annulipes — Bott 1973: 322, fig. 13.

Uca (Celuca) annulipes — Dai & Yang 1991: 467, pl. 3, fig. 362 (2).

Uca (Celuca) lactea annulipes — Crane 1975: 299, 301, 611. figs. 18A–C, 19I –N, 20D–K, 24. — Frith 1977: 6, 13 (in list). — Vannini & Valmori 1981: 208 –210, figs. 5c1–4, 6c.

Lectotype. 1 male (MNHN B11854),"Mer des Indes", leg. Reynaud.

Paralectotype. 1 male (MNHN B11884), data as lectotype.

Other material. Mozambique: 1 male ( ZMH K2913); 2 males ( ZMH K11921 ), 03.08.1927; 1 male ( SMF 17138), Beira, 03.09.1962; 1 male, 1 female ( ZMH K12863 ), E. Africa (possibly Mozambique).

Tanzania: 10 males ( NHMB 576c), leg. La Roche, 1910; 4 males ( SMF 5682), Dar Es Salam, 1899, 1eg. F. Winter; 8 males ( ZSM), 0 9.1890, 1eg. Müller; 3 males, 1 female ( ZMH K29767 ), 10.11.1967; 1 male ( ZMH K2916), Bagamoyo, Lagoon, N. of the city, 26.06.1888, leg. F. Stuhlmann; 1 male ( ZMH K3362), Kingani, near to ferry, mangroves, burrowing in muddy banks, 29.06.1888, leg. F. Stuhlmann; 9 males, 1 female ( ZMH K2916), Kingani, near ferry to Bagamoyo, flooding area, mud, 0 8.07.1888, leg. F. Stuhlmann; 3 males ( ZMH Ex K2993), Zanzibar; 31 males ( ZMH K2898); 5 males ( ZMH K2911); 5 males ( ZSM); 8 males, 6 females ( ZMH K2919), Lagoon, 20.04.1888, 1eg. F. Stuhlmann.

Kenya: 71 males, 2 females (ZSM), Mombasa, leg. Wache; 1 male (ZMH K27111 ), river bank, 02.1957; 1 male (UZM), Mombasa Harbour, Kilindini, mangrove swamp, "Dana" Stat. 394; 1 male (SMF 17139), between Mombasa and Malindi, Kilifi Creek, Manorani, 12.1985, leg. W. Baumeister; 19 males, 7 females (SMF 9107), South of Malindi, Mida Creek, Island, mangroves, 10.1979, 1eg. M. Grasshoff; 5 males, 2 females (MNHN B11873), Gazi, 45 km S. Mombasa, shore, 1911, leg. Alluaud & Jeannel; 2 males, 1 female (SMF 14921), Gazi, mangroves, 30.08.1985, leg. H. Thiel & H. Rumohr; 18 males, 6 females (SMF 35029), S. Malindi, Minda, mangroves, 0 3.08.1989, H. G. Müller.

a, infraorbital region; b, major chela, outer surface; c, major chela, outer surface (regenerated); d, major chela, inner surface; e, merus of major chela, outer surface; f, merus of major chela, inner surface; g, minor chela, outer surface; h, minor chela, outer surface.

Madagascar: 1 male (MNHN B11880); 4 males (MNHN 11889); 1 male (MNHN B11892); 1 male, 1 female (MNHN B11894); 1 male (MNHN B11896); 20 females (MNHN B11904); 4 males (MNHN B11909); 1 male (MNHN B11910); 1 male, 1 female (MNHN B11913); 11 males (MNHN B11900), 1eg. Waterlot; 2 males (MNHN B11899), leg. G. Petit; 11 males, 3 females (SMF 5853), 1880, leg. Stumpf & Ebenau; 1 male (SMF 568l), 1881, leg. A. Stumpf; 3 males, 1 female (MNHN B11890), Nossi Bé; 1 male (MNHN B11879), 6 males (MNHN B11880). 1 male (UZM), leg. A Crosnier; 1 male (MNHN B2024l), id, 11.08.1953, 1eg. T.

Monod; 1 male (ZMH), 1883, leg. A. Stumpf; 2 males (SMF 17142), 0 7.06.1883, leg. A. Stumpf; 1 male, 1 female (MNHN B11893), Tamatave; 2 males, 2 females (MNHN 11898), Majunga Bay, 1926; 13.11.1962, leg. Arnoult; 2 males (MNHN B11902), Marondava, leg. Fauliat; 4 males, 5 females (MNHN B11907), Ampalaza Bay, 1eg. G. Petit; 1 male (MNHN B11906), 2 males (MNHN B11908), Analalava, 1903, 1eg. E. HANTZ; 2 males (1 juv) (MNHN B12085), NW coast, Irohono, mangroves, 21.12.1965, leg. Betsch.

Seychelles: 5 males (SMF 6692), leg. R. Seréne; 14 males, 9 females (SMF 12944), Praslin, freshwater creek about 300 m. from sea, 16.05.1979, 1eg. M. Ackermann; 4 males (ZMH K32308 ), Mahé, 0 6.08.1979, leg. Martens & Gillandt; 2 males (SMF 17143), Port Gland (La Plaine Estate), mangroves, 0 9.09.1977, leg. H. Bosch.

Mauritius: 1 male (MNHN B11875), 1913, leg. P. Carie.

Indian Ocean: 2 males (ZMG 110), leg. Reussmann; 6 males (ZSM), vend. Salmin.

India: 3 males (SMF 9739), vend. Salmin [ex. Mus. Heidelberg]; 17 males, 6 females (ZMH K2977); 5 males (ZMH K26900 ); 27 males, 2 females (ZMH K26913 ); 3 males (SMF 2343), 1 male (ZSM), Bombay, Bandra, shore, 1eg. R. Altevogt; 6 males (ZMH K26870 ), Alibag S. of Bombay, 28.11.1955, leg. G.A. Von Maydell; 11 males (ZMH Ex27091), 0 4.12.1955, leg. G.A. Von Maydell; 7 males, 4 females (UZM) Madras, "Galathea"-Expedition, leg. Reinhardt; 2 males (UZM), 0 4.1908, 1eg. Berg & Mathiassen; 1 male (UZM), Vellore, 1eg. Loeventhal; 1 male (MNHN B11878), Karikal, intertidal, 20.12.1940, 1eg. K. Lindberg; 5 males (UZM), Trankebar, "Galathea"-Expedition; 1 female (MNHN B11853), Vellar estuary near Porto Novo, 14.05.1980, leg. P. Noel; 1 female (MNHN B11881), Pondichery; 17 males, 4 females (MNHN B11888); 3 males, 1 female (ZMK ex Cr.1534), Nankauri.

Sri Lanka: 4 males, 2 females (SMF 7833), Negombo, mangroves, 0 2.04.1977, leg. R. König; 4 males (ZMH K30309 ), Bentota, river estuary; 2 males (ZMH K2912), 0 3.08.1909, 1eg. G. Duncker; 5 males (SMF 5692), Weligama-Bay, 0 4.02.1912, leg. O. Löw-Beer; 7 males (NHMB 576a), Trincomalee, 1886–1893, leg. Sarasin; 6 males (UZM), shore, 0 3.1889, 1eg. K. Ristert.

Thailand: 4 males (UZM), 07.02.1920; 38 males, 3 females (UZM); 6 males, 1 female (UZM), 1915, 1eg. Fogh; 1 male (SMF 9854), Phuket, W. coast, several 100 m. upstream, intertidal zone, 1981, leg. J. Kaden; 1 male (UZM), Terutao Isl., estuary, 01- 02.03.1966 [5th Thai Dan. Exped., Stat. 1149].

Malaysia: 1 male (ZMG 102), W. coast of Malay Peninsula, 1894, leg. Sturm; 5 males (SMF), Malay Peninsula, 1901, 1eg. M. Jensen; 1 male, 3 females (UZM), 3 males (UZM), Telok Merban, shore, 22.08.1937, 1eg. C.W. Frank; 3 males, 2 females (UZM), Pinang, "Galathea"-Expedition, leg. Reinhardt; 2 males, 1 female (ZSM), shore, 17.12.1912, 1eg. Haberer.

Singapore: 1 male (ZSM), 1906, leg. Schauinsland; 1 male (SMF 6685), 2.01.1972, leg. R. Seréne; 6 males (ZMH K3073), 3 males (ZMH K3204), Changi, 10.1900, leg. G. Duncker; 1 male (UZM), Pulau Tekong, south coast, sandy mud flat, 28.05.1951, "Galathea"-Expedition, stat. 351.

Indonesia: 1 female (ZSM), Java, Soetji near Grisee; 7 males (ZSM), 2 males, 1 female (UZM), Tjilatjap; 2 males (NHMB 576b), Celebes, Makassar, 1894, leg. Sarasin.

China: 1 male (IOASQ), Hainan Island, Gangmen, 0 5.12.1955, leg. Liu.

Japan: 1 male (SMF 34768), Ogasawara (Bonin Islands), T. Sakai.

Wrong or doubtful localities: 4 males (ZSM), Chile, 1eg. Wessel; 1 male (ZMH K3205), W. Africa, 1911, leg. W. Michaelsen.

Unknown locality: 25 males, 1 female (MNHN B11865); 2 males (1 Chelip of male) (MNHN B11877); 42 males, 3 females (MNHN B11886); 1 male (MNHN B11895); 2 males (MNHN B11901); 6 males (MNHN B11903); 1 male (MNHN B11905); 1 male (MNHN B11911); 1 male (MNHN B11937); 4 males (UZM); 1 male (ZMH Ex K6748); 1 female (ZMH); 2 males, 5 juv. (SMF 17144).

Diagnosis. Front broad. Anterolateral borders of carapace moderately convergent. Major male chela ( Figs. 2 b–d) with palm bearing a supramarginal groove adjacent to lower border, which very often is beset with short bristles. G1 ( Figs. 3 a–e) with palp relatively short, sometimes reaching to base of horny endpiece ( Figs. 3 b, c); this last one flat, with flanges not inclined, longer than broad, mainly due to the largely protruding dorsal lobe; suture displaced ventrally; terminal opening marked by a more or less distinct notch. Female genital opening as in U. albimana .

Distribution. Indian Ocean: from East Africa to the Indo-Malayan archipelago, excluding the Red Sea and the Persian Gulf, western Pacific: Borneo, Celebes, Lombok, Sumbawa, Flores, Phillippines, Hainan Island of China, probably ranging up to Ogasawara (Bonin) Islands.

Remarks. Crane (1975) treated this species as a subspecies of U. lactea . We do not think that species with such different pleopods do interbreed and therefore assign a species rank to Crane’s subspecies. This is also the case of U. perplexa , U. lactea and U. cryptica sp. nov., which are described herein. In the Persian Gulf populations hitherto referred to as U. annulipes the G1s are within the range of variability of U. annulipes (see Figs. 3 b–e, or Crane 1975: fig. 65D). However, the anterolateral borders of the carapace consistently do not converge in contrast to the convergent anterolateral borders in U. annulipes from outside the Persian Gulf. Furthermore, they have a completely different form of the median tooth of the gastric mill (see Figs. 4 b, 7a). In this respect U. annulipes resembles much more U. albimana , by having 8 teeth on the basal plate of the median tooth, while the Persian Gulf populations have only 6. Therefore we separate these last ones under the name U. iranica Pretzmann, 1971 , which we rank as a morphologically closely related but distinct species. U. albimana ( Kossmann, 1877) and U. mjoebergi Rathbun, 1924 , are easily distinguishable from U. annulipes by lacking the supramarginal groove of the large cheliped. Regarding the form of the median tooth of the gastric mill ( Fig. 4 b), U. annulipes is closely related to U. albimana rather than to U. mjoebergi , the latter having a fully different morphology in the median tooth. Therefore these two taxa are treated as full species here. Shih et al. (2009), based on genetic evidence, identified two species, U. albimana and U. annulipes as good species, and assigned them together with U. iranica in the same clade, which can be well supported by morphological findings.

Small variations on the apical portion of the G1 of U. annulipes are observed in specimens from different localities (see Figs. 3 b–e), but these differences are not considerable as all have the same pattern by having a short palp not reaching to the base of endpiece, a long apical flange, distally two lobed, and in all cases the dorsal lobe is larger than ventral one. There is another noteworthy feature not noticed in earlier studies. The gap of the small chela in males is always narrower than the breadth of the adjacent dactylus in all specimens from Thailand, farther east than India ( Fig. 2 h). In East African samples, a certain number of specimens have a very broad gap ( Fig. 2 g), being superior to the breadth of the adjacent dactylus. However, this character is not consistent in that particular area. This probably can be the beginning of a differentiation of western and eastern Indian Ocean forms that may be taking place, as the character is asymmetrically distributed. Because of the missing consistency we do not advocate any taxonomic separation at the present time. As may be expected, the identification of females of wide-front Uca related to U. annulipes is very difficult and seems impossible in most forms. We have examined the small chelae of females, which showed no differences between different species, as well as the female genital opening. Of all the species, only U. perplexa is slightly different by the orientation of the genital opening.

Figures

FIGURE 2. Uca annulipes (H. Milne Edwards, 1837): a, male (SMF 14921), Kenya; b, male (SMF), Malay Peninsula; c, male (SMF 6692), Seychelles; d – f, male (ZSM), Kenya, Mombasa; g, male (SMF 5682), Tanzania, Dar Es Salam; h, male (SMF 9854), Thailand.

FIGURE 3. Uca annulipes (H. Milne Edwards, 1837): a – c, male (SMF 12944), Seychelles; d, male (ZSM), Kenya, Dar Es Salam; e, male (SMF 9739) India. a, G 1, lateral surface; b, apical part of G 1, lateral surface; c, apical part of G 1, mesial surface; d, apical part of G 1, lateral surface; e, apical part of G 1, lateral surface.

FIGURE 4. Median tooth plate of the gastric mill, ventral face. a, Uca albimana (Kossmann, 1877), male (SMF 26018); b, Uca annulipes (H. Milne Edwards, 1837), male (SMF 9107).