Search

Spirobranchus sinuspersicus sp. nov. (Figs 2, 3 A–D, 4, 5A–D, 6) Synonymy. Spirobranchus maldivensis. —not Pixell, 1913 but sensu Wesenberg-Lund (1949): 358–359 [Jask, Gulf of Oman, Iran, diagno- sis]. Pomatoleios kraussii.— Mohammad (1971: 300) [Kuwait, short description of tubes]; Mohammad (1975: 1–15) [Kuwait, ecol- ogy]; Crisp (1977: 147–160) [Kuwait, larval development]; Fiege (1992: 1–23) [Saudi Arabia, “even found on heavily oiled rocks/beaches”]; Wehe & Fiege (2002: 129) [name only, checklist of Persian Gulf polychaetes], Al-Yamani et al. (2012: 75) [Kuwait, name and photos of tubes]; Joydas et al. (2012: 330) [PG, Saudi Arabia, name only]; Safari et al. (2014a, b) [Bandar Abbas, Strait of Hormuz, Iran, description]. Spirobranchus kraussii.— Pillai (2009: 168, Fig. 49E–G) [Abu Dhabi, United Arab Emirates, short description]; Baghernejhad (2012: 70) [Bushehr, PG, Iran, name and abundance], Baghernejhad et al. (2015: 71, 73) [Bushehr, PG, Iran, population dynamics]; Lavajoo & Amrollahi-Biuki (2015: 49–57) [Bandar Abbas, Strait of Hormuz, Iran, reproductive ecology]; Bonyadi-Naeini et al. (2017: 2–6) [Abu-Musa Island, PG, Iran, name only]; Sánchez Ovando (2019: 45, Fig. 14A–C, 23) [Dubai, the United Arab Emirates; description]; Lavajoo (2019: 93–97) [Bandar Abbas, Strait of Hormuz, Iran, larval ecol- ogy]; Shabani et al. (2019: 1–9) [Northern coast of the Persian Gulf, ecology]. Spirobranchus cf. kraussii .— Al-Kandari et al. (2019: 8) [Kuwait, PG, name only, checklist of intertidal polychaetes of Ku- wait]. Galeolaria sp.— Roohi-Shalmaee et al. (2019) [Bandar Abbas, Strait of Hormuz, Iran, ecology]. Type material. Holotype: St.7, ZUTC.6780, on rock, high intertidal zone, legit S. Pazoki, 4A–B, Table 1. Paratypes: total of 28 specimens, Fig. 1A, Table 1. Persian Gulf: St.1, ZUTC.6781 (1 spec.), ZUTC.6783 (1 spec.), ZUTC.6807 (1 spec.); St.2, ZUTC.6800 (1 juvenile), ZUTC.6806 (1 spec.); St.6, ZUTC.6779 (1 spec.); St.7, ZUTC.6782 (1 spec.), ZUTC.6805 (1 spec.); St.9, ZUTC.6784 (1 spec.), NHMO C7015 (1 spec.), ZUTC.6786 (1 spec.), ZUTC.6801 (1 juvenile); Strait of Hormuz: St.13, ZUTC.6796 (1 spec.), ZUTC.6797 (1 spec.); St.14, ZUTC.6791 (1 spec.); St.15, ZUTC.6788 (1 spec.), ZUTC.6789 (1 spec.), ZUTC.6799 (1 juvenile), ZUTC.6812 (1 juvenile), ZUTC.6813 (1 juvenile); St.17, ZUTC.6778 (1 spec.), NHMO C7016 (1 spec.), ZUTC.6802 (1 juvenile), ZUTC.6808 (1 spec.); St.18, AM W.52757 (1 spec.), ZUTC.6804 (1 spec. prepared for SEM); Gulf of Oman: St.22, ZUTC.6790 (1 spec.), AM W.52758 (1 spec.), legit. H. Ashrafi; St.25, ZUTC.6793 (1 spec.), ZUTC.6794 (1 spec.), ZUTC.6795 (1 spec.), ZUTC.6809 (1 spec.), ZUTC.6810 (1 spec.); all material legit by S. Pazoki from mid to high intertidal unless otherwise stated. Material not included in the type series. Specimens with fringed peduncular wings: St.7, ZUTC.6815 (1 spec.); St.18, ZUTC.6816 (1 spec.); St.25, ZUTC.6814 (1 spec.); specimens with dissected talons: St.6, ZUTC. 6819 (1 spec.); St.14, ZUTC. 6820 (5 spec.); St.15, ZUTC.6818 (1 spec.), ZUTC. 6821 (7 spec.), St.17, ZUTC. 6822 (7 spec.), St.23, ZUTC. 6822 (7 spec.); St.25, ZUTC.6817 (1 spec.). Material used for comparison. Spirobranchus kraussii, Danger Point, Walker Bay, South Africa, 34°29’S, 19°21’E, 10 Apr 2017, ZUTC.6803, legit H. van Niekerk (5 spec., one with opercular endplate dissected). Description. Measurements are shown in the format of “mean ± SD [range/ holotype]” based on a holotype and 20 paratypes. The terminology follows that of ten Hove & Kupriyanova (2009). Tube trapezoidal or sub-triangular in cross section, 3.6 ± 0.8 mm [2.1–4.9/4.8] wide with a lumen of 1.8 ± 0.5 [0.9–3.0/2.5] mm in diameter and a projection from two longitudinal ridges expanding anteriorly making a flap over tube mouth (Fig. 2C, D). In adults, flap expanded more laterally and anteriorly. However, juvenile tubes sometimes with a prominent wavy keel (Fig. 2 A–B). Outer surface of tube white, pale blue or violet (sometimes pink in small juveniles), inner surface white, blue or violet. Radiolar crown composed of two radiolar lobes, each lobe with a mean of 16.6 ± 3.4 [12–20/20] radioles ar- ranged in a semicircle. Inter-radiolar membrane extending up to 48% ± 13% [30–66/59%] of radiolar length, usu- ally white, making a whitish band between the basal half of crown and free end of radioles. Prostomial eyes absent. Radiolar eyes in three to six pairs per radiole (Fig. 4D). Stylodes absent. Radiolar crown violet, greenish blue, red, pink, yellow, and white (Fig. 3 A–D). Juveniles sometimes with lighter colours. Peduncle triangular in cross-section, inserted near medial line between radiolar lobes (Fig. 4C), distal part dorso-ventrally flattened; usually pigmented with dark and light bands, a total length of 1.9 ± 0.5 mm (1.1–2.6/ 1.8 mm). A transverse line making a two-segmented appearance of peduncle, 44.8 ± 6.2% (35–54/47%, Fig. 4 B–C) from the proximal part of the peduncle. Peduncular wings with a smooth edge and a tapering tip (Fig. 4 B–C), but rarely fringed with 2–5 finger-like processes (seven specimens) from Dayyer (ZUTC.6815), Ramin (ZUTC.6814), and Hormuz (ZUTC.6816, Fig. 4D). Unpaired proximal wing absent. Pseudoperculum absent. Operculum made of massive flattened ampulla covered with a white calcareous circular endplate (Fig. 4 A–D), 1.6 ± 0.4 mm (1.0–2.3/ 1.7 mm) in diameter, somewhat concave with a non-transparent rim, without any processes distally (Figs 4 C–D, 5A). Talon proximally inserted into tissue of opercular ampulla, thus usually not visible without dissection or clearing up tissues; talon consisting of two symmetrical dorsal or three arranged in a triangle dorsoventral protuberances, with a depression in between, where endplate meets tissue (Fig. 5 B–D). Collar and thoracic membranes. Collar well-developed, sometimes as long as operculum (Fig. 3A, C), with entire edge; trilobed, ventral lobe broader than dorso-lateral ones. Dorso-lateral lobes continuing in thoracic membranes forming an apron over first or second abdominal chaetigers. A pair of triangular tonguelets between lateral and ventral lobes present (Fig. 4E). Collar creamy to dark blue or violet. Thorax with six thoracic uncinigerous segments; collar chaetae absent in adults, but present in juveniles (Fig. 4F). Thoracic chaetae limbate of two sizes (Fig. 6C). Uncini saw-shaped with 7–10 teeth and flat gouged peg (Fig. 6A). Thoracic tori similar in size along thorax, widely separated anteriorly, and gradually approaching posteriorly, forming a ventral depression (Fig. 3 B–D). Abdomen. One or two achaetous segments anteriorly, number of chaetigers 41 ± 6 [31–50/38]. Abdominal chae- tae with long shaft present throughout abdomen (Fig. 6E); distal end true trumpet-shaped, one side drawn out into a tapering process (Fig. 6F). Uncini saw-shaped, similar to thoracic ones, with 8–12 teeth (Fig. 6B, D) from anterior to posterior abdomen. Chaetae becoming increasingly longer posteriorly, but capillary chaeta absent in posterior chaetigers. Adults body proper brownish, yellow, red, green, but predominantly blue to violet (Fig. 3). Size. Total length 8.6 ± 2.7 mm [4.8–15.4/ 8.8 mm]; width of thorax 1.8 ± 0.5 mm [1.0–3.3/ 1.6 mm]; radioles accounting for 22.0 ± 5.1% [11.9–29.4%] of entire length. Habitat. Animals build large dense tube aggregations of 600,000 ± 15,000 specimens/m 2, with a maximum density occurring in summer (Safari et al. 2014b) in the upper intertidal zone, attached to the upper surface of rocks (Fig. 2 F–G) or any hard substrates, including man-made structures, such as breakwater walls, concrete piers and rock jetties (Table 2). Type locality. Dayyer, Bushehr Province, Persian Gulf, Iran, 27°50’4.688”N, 51°57’0.143”E. Geographic distribution. Persian Gulf: coasts of Iran, Kuwait, Saudi Arabia, the United Arab Emirates (Abu Dhabi, Dubai); Gulf of Oman: Iran. Reproduction and development. According to Crisp (1977), egg size in Spirobranchus sinuspersicus sp. nov. (as Pomatoleios kraussii) is 60–65 µm and larvae show planktotrophic development typical for the genus (Kupriyanova et al. 2001). Larvae complete their development to metamorphosis in 1–3 weeks at 25–27°C being fed by micro- flagellate Isochrysis galbana Parke or Tetraselmis suecica (Kylin) Butcher cultures under laboratory conditions. Competent larvae settle and metamorphose preferentially on the tubes of adults, especially on newly-settled ones (Crisp 1977). In the field, larval settlement takes place from March to December with a maximum in August, which corresponds to water temperature of 30°C (Mohammad 1975). Taxonomic remarks. Specimens of Spirobranchus sinuspersicus sp. nov. described herein and S. kraussii from the type locality in South Africa are superficially similar in most of their morphological characters, such as the lack of collar chaetae in adults, shape of the tube and unadorned opercular endplate, as well in being gregarious and living in intertidal habitats. Nevertheless, these two species are morphologically recognizable. The largest specimens of Spirobranchus sinuspersicus sp. nov. are noticeably smaller than those of S. kraussii 15 vs. 31 mm, respectively (Fig. 3E). Differences in size are evident even in juvenile specimens (2.5–3.5 mm vs. 9.6–11.7 mm, respectively). Abdominal chaetigers are more numerous in S. kraussii than in S. sinuspersicus sp. nov. (70 ± 10 and 41 ± 6 segments, respectively). Pillai (2009), who had studied specimens of S. sinuspersicus sp. nov. (identified as Pomatoleios kraussii) from Abu Dhabi, has reported the length of adult specimens to be 23.0 mm. However, among hundreds of specimens collected in the present study, none was even close to that size. Although Pillai (2009) did not mention the number of abdominal segments, based on his figure (Pillai 2009, Fig. 49E, F), there are around 30 chaetigers. Probably the most important morphological difference between Spirobranchus sinuspersicus sp. nov. and S. kraussii is the shape of the talon; a bulge made of two or three protuberances and a concavity between them in S. sinuspersicus sp. nov. (Fig. 5 A–D) and as an extension made of about 10 protrusions in S. kraussii (Fig. 5 E–F). This feature has been mentioned in specimens from Abu Dhabi (Pillai 2009), but reported for 16 out of 18 specimens from Kuwait by ten Hove (Simon et al. 2019). The major problem with this character is that its variability is not known. Spirobranchus sinuspersicus sp. nov. also differs from S. kraussii by some more subtle characters. In S. sinuspersicus sp. nov. peduncular wings originate from the base of the peduncle resulting in a “V” shape appearance (Fig. 4 B–C), while in S. kraussii the wings originate from the mid-peduncle, resulting in a “Y” shaped appearance (Fig. 3G). Furthermore, the opercular peduncle is located centrally between the radiolar lobes in S. sinuspersicus sp. nov. (Fig. 4 B–C), but it is inserted below left radiolar lobe in S. kraussii (Fig. 3F; Simon et al. 2019). The abdominal chaetae in S. sinuspersicus sp. nov. are present from second or third abdominal segments, while they are present only in about 10 posterior abdominal segments of S. kraussii specimens. The presence of fringed peduncular wings described herein have not been reported either for Spirobranchus kraussii or other for any species of the S. kraussii -complex, but such occasional presence might easily have been overlooked. Fringed wings are common for Spirobranchus species such as S. cf. tetraceros (Ben-Eliahu & ten Hove 2011: 88, 93, Table 5) and S. cf. polytrema (Phillippi, 1844) and populations of these taxa also have specimens with simple unfringed distal wings (ten Hove, pers. comm.). However, in S. sinuspersicus sp. nov., the wings are normally smooth, rarely fringed. Variability of this character, however, has been documented insufficiently in the literature. Our morphological study also revealed the presence of occasional saw to rasp-shaped uncini in the new species. Some thoracic and abdominal uncini may have 2 or 3 teeth in one or two row(s) above peg in Spirobranchus sinuspersicus sp. nov. (Fig. 6B), which was also seen in S. kraussii (unpublished SEM images by C. Simon, pers. comm.). Such uncini are also found in other species of Spirobranchus (ten Hove & Kupriyanova 2009: 96). Presence of occasional two teeth in the middle of a typical saw-shaped uncini (Fig. 6D) is an unusual feature of S. sinuspersicus sp. nov. Additionally, the basal part of the posteriormost tooth of both thoracic and abdominal uncini maybe split in both S. sinuspersicus sp. nov. and S. kraussii (Fig. 6B and Simon’s unpublished photos). Ecological observations. Spirobranchus kraussii inhabits the high to low intertidal zone, depending on wave exposure rate (Simon et al. 2019). Aggregations of S. sinuspersicus sp. nov. were most abundant in high intertidal zones of sheltered to moderately exposed cliffs, on pebbles and rocks. They were hardly ever found in sub-tidal habitats, and if so, never in aggregations. Our observations are consistent with those of Crisp (1977) who reported preference of this species to settle on natural rather than artificial substrates. However, Mohammad (1975) reported S. sinuspersicus sp. nov. (as Pomatoleios kraussii) from experimental settlement plates made of teak wood and some specimens were recently observed on PVC panels suspended 1 m deep for 3, 6 and 9 months in five Iranian ports (Table 2, St. 3, 8, 10, 11, 12, A. Nasrollahi, unpubl. data). Large aggregations were found on man-made structures, such as breakwater walls made of large stones (Fig. 2G) and on cement wharf pilings and rock jetties in ports of Dayyer, Bushehr, Kong, and Jask. Shabani et al. (2019) reported the species (as S. kraussii) on anthropogenic marine debris (e.g., plastic bottles, glass jars and wood) stranded on the coasts of the PG, arguing that this species can attach to various floating items regardless of the material type and surface texture. However, we did not find specimens of the new species either on floating substrates (such as buoys and ropes) or on hulls of small vessels such as ferries, fishing boats (open dories), and traditional dhows. Spirobranchus sinuspersicus sp. nov. co-occurs with intertidal barnacles Amphibalanus amphitrite (Darwin, 1854) and Chthamalus barnesi Achituv & Safriel, 1980, as well as rock oysters Saccostrea cuccullata (Born, 1778). Spirobranchus sinuspersicus sp. nov. and C. barnesi make a distinct zone in the high intertidal, while A. amphitrite and S. cuccullata dominate lower intertidal where wave action is considerably strong (R. Naderloo, pers. comm.). According to Mohammad (1975), the bathymetric distribution of the worm extended between 0.4 and 1.9 m above datum (level below which the tide seldom falls) where specimens of S. sinuspersicus sp. nov. are limited by other encrusting species (e.g. ectoprocts, sponges and filamentous algae). In artificial bays, ports and on piers, S. sinuspersicus sp. nov. was relatively more abundant than either of the other three species. Etymology. The specific epithet is an adjective and refers to the Persian Gulf (Sinus Persicus in Latin), the type locality of the new species., Published as part of Pazoki, Samaneh, Rahimian, Hassan, Struck, Torsten H., Katouzian, Ahmad R. & Kupriyanova, Elena K., 2020, A new species of the Spirobranchus kraussii - complex (Annelida, Serpulidae) from the Persian Gulf and Gulf of Oman, pp. 401-430 in Zootaxa 4748 (3) on pages 406-414, DOI: 10.11646/zootaxa.4748.3.1, http://zenodo.org/record/3700975, {"references":["Pixell, H. L. M. (1913) Polychaeta of the Indian Ocean, together with some species from the Cape Verde Islands. Transactions of the Linnean Society of London, Zoology, 16, 69 - 92. https: // doi. org / 10.1111 / j. 1096 - 3642.1914. tb 00125. x","Wesenberg-Lund, E. (1949) Polychaetes of the Iranian Gulf. In: Jessen, K. & Sparck, R. (Eds.), Danish Scientific Investigations in Iran. Vol. 4. Ejnar Munskgaard, Copenhagen, pp. 247 - 400.","Mohammad, M. B. (1971) Intertidal polychaetes from Kuwait, Arabian Gulf, with descriptions of three new species. Journal of Zoology, 163 (3), 285 - 303. https: // doi. org / 10.1111 / j. 1469 - 7998.1971. tb 04536. x","Mohammad, M. B. (1975) Competitive relationship between Balanus amphitrite amphitrite and Pomatoleios kraussii with special reference to their larval settlement. Hydrobiologia, 46 (1), 1 - 15. https: // doi. org / 10.1007 / BF 00038722","Crisp, M. (1977) The development of the serpulid Pomatoleios kraussii (Annelida, Polychaeta). Journal of Zoology, 183 (2), 147 - 160. https: // doi. org / 10.1111 / j. 1469 - 7998.1977. tb 04178. x","Fiege, D. (1992) Polychaeta of the Intertidal Zone in the Ras Az-Zawr Marduma Bay Region, Saudi Arabia. Unpublished Progress Report of the Senckenbergische Forschungsinstitut, Frankfurt am Main, 23 pp.","Wehe, T. & Fiege, D. (2002) Annotated checklist of the polychaete species of the seas surrounding the Arabian Peninsula: Red Sea, Gulf of Aden, Arabian Sea, Gulf of Oman, Arabian Gulf. Fauna of Arabia, 19, 7 - 238.","Al-Yamani, F. Y., Skryabin, V., Boltachova, N., Revkov, N., Makarov, M., Grintsov, V. & Kolesnikova, E. (2012) Illustrated Atlas on the Zoobenthos of Kuwait. Kuwait Institute for Scientific Research, Kuwait City, 383 pp. http: // doi. org / 10.13140 / RG. 2.1.2535.7285","Joydas, T. V., Qurban, M. A., Al-Suwailem, A., Krishnakumar, P. K., Nazeer, Z. & Cali, N. A. (2012) Macrobenthic community structure in the northern Saudi waters of the Gulf, 14 years after the 1991 oil spill. Marine Pollution Bulletin, 64 (2), 325 - 335. https: // doi. org / 10.1016 / j. marpolbul. 2011.11.007","Safari, N., Amrollahi, N., Kamrani, E. & Shahdadi, A. (2014 a) Identification of tube worms Serpulidae of Shilat Bay in Bandar Abbas. First regional conference on the sea, the development and water resources of coastal areas of Persian Gulf, Bandar Abbass, Hormuzgan, No. 34, 1 - 10. [in Persian, https: // www. civilica. com / Paper-WATERJDH 01 - WATERJDH 01 _ 034. html]","Pillai, T. (2009) Descriptions of new serpulid polychaetes from the Kimberleys of Australia and discussion of Australian and Indo-West Pacific species of Spirobranchus and superficially similar taxa. Records of the Australian Museum, 61 (2), 93 - 199. https: // doi. org / 10.3853 / j. 0067 - 1975.61.2009.1489","Baghernejhad, N. (2012) Survey on seasonal changes on biodiversity, richness and distribution of intertidal polychaetes of Bushehr rocky shores. Master's thesis. University of Bushehr, Bushehr, 77 pp. [in Persian]","Baghernejhad, N., Salari, M. A., Ronagh, M. T. & Vazirizadeh, A. (2015) The effect of seasonal variations on diversity and dominance of intertidal Polychaeta of the Persian Gulf (Bushehr Province). Journal of Oceanography, 6 (22), 69 - 75. [in Persian, http: // joc. inio. ac. ir / article- 1 - 771 - fa. html]","Lavajoo, F. & Amrollahi-Biuki, N. (2015) Biological structure and the effect of temperature on embryonic stage of the Spirobranchus kraussii (Baird, 1865) (Annelida: Serpulidae) in Bandar Abbas in the Persian Gulf. Journal of the Persian Gulf, 6 (22), 49 - 60. [http: // jpg. inio. ac. ir / article- 1 - 445 - en. html]","Sanchez Ovando, J. P. (2019) Revision de las especies del genero Spirobranchus Blainville, 1818 (Polychaeta: Serpulidae) de America tropical. Bachelor's thesis, Universidad del Mar, Puerto Angel, Oaxaca, Mexico, 99 pp.","Lavajoo, F. (2019) Influence of different algal diets on larval growth rates in the marine Serpulidae polychaete worm Spirobranchus kraussii. Croatian Journal of Fisheries, 77 (2), 93 - 98. https: // doi. org / 10.2478 / cjf- 2019 - 0010","Shabani, F., Nasrolahi, A. & Thiel, M. (2019) Assemblage of encrusting organisms on floating anthropogenic debris along the northern coast of the Persian Gulf. Environmental Pollution, 254, 112979. https: // doi. org / 10.1016 / j. envpol. 2019.112979","Al-Kandari, M., Sattari, Z., Hussain, S., Radashevsky, V. & Zhadan, A. (2019) Checklist of intertidal polychaetes (Annelida) of Kuwait, Northern part of the Arabian Gulf. Regional Studies in Marine Science, 2019, 100872. https: // doi. org / 10.1016 / j. rsma. 2019.100872","Roohi-Shalmaee, N., Mousavi-Nadushan, R., Mostafavi, P. G., Shahbazzadeh, D. & Bagheri, K. P. (2019) Ecological adaptation of the Persian Gulf polychaete in a polluted area: proteomics concerning dominant defensive biomarkers. International Journal of Environmental Science and Technology, 2019, 1 - 10. https: // doi. org / 10.1007 / s 13762 - 019 - 02508 - y","Kupriyanova, E. K., ten Hove, H. A., Sket, B., Zaksek, V., Trontelj, P. & Rouse, G. W. (2009) Evolution of the unique freshwater cave-dwelling tube worm Marifugia cavatica (Annelida: Serpulidae). Systematics and Biodiversity, 7 (4), 389 - 401. https: // doi. org / 10.1017 / S 1477200009990168","Safari, N., Amrollahi, N., Kamrani, E., Shahdadi, A. & Razaghi, M. (2014 b) A survey on eco-biological traits of Pomatoleios kraussii of Bandar Abbas, Shilat Bay. First regional conference on the sea, the development and water resources of coastal areas of Persian Gulf, Bandar Abbass, Hormuzgan, No. 34, 1 - 10. [in Persian] [https: // www. civilica. com / Paper-WATER- JDH 01 - WATERJDH 01 _ 037. html]","Kupriyanova, E. K., Nishi, E., Hove, H. A. ten & Rzhavsky, A. V. (2001) A review of life-history patterns in serpulimorph polychaetes: Ecological and evolutionary perspectives. Oceanography and Marine Biology: an Annual Review, 39, 1 - 100.","Simon, C. A., van Niekerk, H. H., Burghardt, I., ten Hove, H. A. & Kupriyanova, E. (2019) Not out of Africa: Spirobranchus kraussii (Baird, 1865) is not a global fouling and invasive serpulid of Indo-Pacific origin. Aquatic Invasions, 14 (2), 221 - 249. https: // doi. org / 10.3391 / ai. 2019.14.2.05","Ben-Eliahu, M. N. & Hove, H. A. ten (2011) Serpulidae (Annelida: Polychaeta) from the Suez Canal-from a Lessepsian migration perspective (a monograph). Zootaxa, 2848, 1 - 147. https: // doi. org / 10.11646 / zootaxa. 2848.1.1","Phillippi, A. (1844) Einige Bemerkungen uber die Gattung Serpula, nebst Aufzahlung der von mir im Mittelmeer mit dem Their beobachteten Arten. Archiv fur Naturgeschichte, Berlin, 10 (1), 186 - 198. https: // doi. org / 10.5962 / bhl. part. 29558","Darwin, C. (1854) A Monograph on the Sub-Class Cirripedia with Figures of All the Species. The Balanidae, (or Sessile Cirripedia); the Verricidae, etc., etc., etc. The Ray Society, London, viii + 684 pp.","Achituv, Y. & Safriel, U. N. (1980) A new Chthamalus (Crustacea: Cirripedia) from intertidal rocks of the Red Sea. Israel Journal of Ecology and Evolution, 29 (1 - 3), 99 - 109. http: // doi. org / 10.1080 / 00212210.1980.10688487","Born, I. von (1778) Index Rerum Naturalium Musei Caesarei Vindobonensis. Pars I. ma. Testacea. Verzeichniss der naturlichen Seltenheiten des k. k. Naturalien Cabinets zu Wien. Erster Theil. Schalthiere. Kraus, Vindobonae, XL + 458 pp. https: // doi. org / 10.5962 / bhl. title. 11581"]}

Tuerkayana rotundum (Quoy & Gaimard, 1824) (Figs. 1, 2, 5, 6, 7A, B, 8A–E, 9A–C, 10A–C, E–G, 11A–C, E–G, I–K) Thelphusa rotunda Quoy & Gaimard, 1824: 527, pl. 77 fig. 1 [type locality: as “Océanie” = Guam]; NK Ng et al. 2019: 101, fig. 3B [lectotype designation]. Cardisoma frontalis H. Milne-Edwards, 1853: 204; De Man 1902: 548, pl. 20 fig. 14; Tweedie 1950: 138, fig. 4h. Discoplax pagenstecheri Kossmann, 1878: 255. Cardisoma rotundum – Türkay 1974a: 234, figs. 1, 14 (part); Sakai 1976: 679, pl. 232; Türkay and K. Sakai 1976: 14, figs. 3, 5, pl. 2(1–2); Takeda 1982: 225; Miyake 1983: 188, pl. 63(3); Hirata et al. 1988: 5, 1 unnumbered fig.; Ho et al. 1992: 61, fig. 6; Poupin 1994: fig. 50; PKL Ng 1998: 1151, 1 unnumbered fig.; Liu 1999: 85, figs. 30–31. Discoplax rotundum – Lee 2001: 140, 2 unnumbered figs.; PKL Ng et al. 2001: 49; Davie 2002: 185; Lin et al. 2011: 37, 2 unnumbered figs.; PKL Ng et al. 2017: 97. Discoplax rotunda – Innocenti and Vannini 2007: 635, fig. 1; PKL Ng et al. 2008: 214; Poupin and Juncker 2008: 35, fig. 5d; Poupin 2010: 42 (part); Poupin and Juncker 2010: 56 (part), 1 unnumbered fig. on p. 57; Orchard 2012: 149, 6 unnumbered figs.; Li and Chiu 2013: 65, 3 unnumbered figs.; Shih 2013 b: 28, fig. 9; PKL Ng and Shih 2014: 111; Toyota et al. 2014: 180, 7 unnumbered figs.; PKL Ng and Shih 2015: 382; Li and Chiu 2019a: 98, 3 unnumbered figs.; Li and Chiu 2019b: 2 unnumbered figs. on p. 52. Tuerkayana rotundum – Guinot et al. 2018: 551, fig. 4G; Sasaki 2019: 12702; Toyota et al. 2019: 188, 4 unnumbered figs. Diagnosis: Frontal, epigastric, epibranchial and anterior part of mesobranchial regions covered with numerous small granules and striae, those on anterior part more flattened; mesogastric, metagastric regions almost smooth; posterior part of meso-, metabranchial regions with prominent oblique striae and granules; epigastric regions prominent, margin relatively sharp; postorbital cristae strong, separated from epigastric by shallow concavity; external orbital tooth broadly triangular, margin confluent with anterolateral margin, separated by short narrow fissure or small cleft (Figs. 2, 5, 6, 7B). Merus of third maxilliped subovate, longer than broad (Fig. 7A). Surfaces of adult chelae with low granules on margins, punctate to rugose (Fig. 9A–C). Ambulatory legs short; ratios of maximum length to maximum width of third and fourth ambulatory meri 2.5–2.6, 2.5–2.6, respectively; surfaces covered with striae, appears rugose; dorsal margin of merus rugose, uneven; lateral margins of propodus, dactylus lined with strong, stiff spines or setae (Figs. 2, 10A–C, E–G). Male pleon relatively broad; somite 6 wider than long, lateral margins gently to strongly convex (Fig. 8A–F). Adult G1 almost straight, when viewed from ventral perspective; distal part curved at angle of about 45° from vertical; distal part of outer margin forming gentle shelf–like structure; distal part gently curves towards corneous tip, basal part tapers gradually to truncate tip in dorsomesial view, outer margin appearing gently concave in ventral view, forming relatively narrow triangular flange (Fig. 11A–C, E–G, I–K). Eastern Indian Ocean to Niue. © 2023 Academia Sinica, Taiwan © 2023 Academia Sinica, Taiwan Material examined: Guam – lectotype male (63.0 × 50.0 mm) (MNHN-IU-2000-3745 = MNHN-B3745) [photographs examined], Guam, Oceania, M. Guérin; 1 male (ZRC 2000.568), coll. P.K.L. Ng, October 1996; 1 female (ZRC 2000.567), Hawaiian Rock Quarry, central Guam, coll. 20 April 2000; 2 females (ZRC 2001.703), Merizo, Guam, coll. P.K.L. Ng, 1 August 2001; 1 male (50.2 × 38.6 mm), 1 female (48.2 × 36.9 mm, with hatched zoeae) (ZRC 2000.566), Pago Bay, outside of marine laboratory, University of Guam, coll. P.K.L. Ng and C.-H. Wang, 15-18 April 2000. Taiwan – 1 male (ZRC 1998.471), Pingtung County, coll. H.-C. Liu et al., 13 May 1998; 1 male (53.3 × 41.0 mm) (ZRC 1998.474), 1 male, 1 female (ZRC 1998.432), Pingtung County, Taiwan, coll. H.- C. Liu et al., 22 May 1998; 1 female (ZRC 1998.476), Pingtung County, Taiwan, coll. H.-C. Liu et al., 18 Sep. 1997; 1 male (37.5 × 29.3 mm) (NCHUZOOL 15886), Siangjiaowan, Pingtung County, coll. 2 September 2000; 1 female (39.4 × 31.5 mm) (NCHUZOOL 15887), Dingtanzih, Kenting, Pingtung County, coll. 17 March 2018; 1 ovig. female (33.8 × 25.0 mm) (NCHUZOOL 15888), Dingtanzih, Kenting, Pingtung County, 27 August 2018. Japan – 1 male (SMF 6382), south of Daito-Jima, Riu-Kiu (= Ryukyu Islands), no other data. Philippines – 1 female (ZRC 2001.312), Tawala Cave, Panglao Island, Bohol, Philippines, coll. P.K.L. Ng et al., 17 December 2000; 1 male (47.8 × 37.8 mm) (ZRC 2008.506), Virata Cave, Panglao Island, Bohol, coll. Manong et al., 26 November 2001; 2 males (ZRC 2008.1005), Hinagdanan Cave, northern part of Panglao Island, Bohol, coll. local collectors, June 2002; 1 male (ZRC 2001.2311), Baloc-Baloc Cave, near Virata Cave, Panglao Island, Bohol, coll. Manong et al., 30 November 2001; 1 male (ZRC 2014.463), Panglao Island Nature Resort, Hinagdanan Cave, coll. P.K.L. Ng, 3 March 2004; 1 male (ZRC 2021.865), station M59, Cambagat Cave, Panglao, 9°37.5'N 123°48.1'E, coll. PANGLAO 2004 Expedition, 5 July 2004; 3 males, 3 females (ZRC 2012.437), outside cave at Panglao Island Nature Resort, Panglao Island, Bohol, coll. P.K.L. Ng and P.Y.C. Ng, December 2010; 1 female (ZRC 2001.2312), Balicasag Island, Panglao Island, Bohol, A. Porpetcho and R. Diaz, 30 November 2001; 1 male (SMF 19745), east Mactan, northwest coast of Hilutangan Channel, Cebu, 10°17'N 124°00'E, coll. A. Schreiber. Indonesia – 1 male (33.9 × 26.9 mm) (SMF), station Ip-1, Ambon, coll. Rumphius Expedition, K. Moosa, 17 January 1973; 1 ovigerous female (ZRC 2010.423), Kri Island, West Papua, coll. H.H. Tan, December 2010. Papua New Guinea – 1 female (SMF 24451), Kavieng, New Ireland, coll. 19 July 1998. Admiralty Islands – 2 males (36.6 × 28.8 mm, 35.9 × 28.3 mm), 1 female (40.4 × 30.9 mm) (SMF 5854), Nissan Atoll, Pinipal, Hanseatischen Südsee-Expedition, coll. E. Wolff, 8 May 1909; 1 female (53.1 × 41.0 mm) (SMF 5855), Pak, Hanseatischen Südsee-Expedition, coll. E. Wolff, 10 September 1909. New Caledonia – 1 male (61.4 × 78.0 mm) (MNHN–B 24813), Easo Cave, 17 m depth, Lifou Island, Loyalty Islands, coll. B. Richer de Forges, 20 August 1993; 1 male (28.0 × 26.3 mm) (MNHN–B 24809), on the ground, Lifou Island, Loyalty Islands, coll. B. Richer de Forges, 18 April 1993; 2 females (38.0 × 48.0 mm, 39.0 × 49.0 mm) (MNHN-B 24814), sea level, Maré Island, Loyalty Islands, coll. B. Richer de Forges, 16 July 1993; 1 juvenile (3.0 × 3.5 mm) (MNHN–B 26947), cave, station 88–066, Ouvea Atoll, Loyalty Islands, coll. T.M. Iliffe and S. Sarbu, 22 June 1988; 9 juveniles (MNHN-B 26948), Loyalty Islands, Luecilla Cave, station 88–058, Lifou Island, coll. T.M. Iliffe and S. Sarbu, 17 June 1988; 3 males (47.2 × 37.6, 47.2 × 37.7, 52.9 × 41.9 mm) (MNHN-IU-2014-10106), Lifou I. (Hnyekej), coll. J. Poupin and M. Juncker, 2005; 1 male (36.0 × 44.0 mm) (MNHN-B 17134), no other data. Solomon Islands – 1 young female, 6 juveniles (12.7 × 16.5 mm) (MNHN-B 26943), Malaita, coll. T.M. Iliffe and S. Sarbu, 27 August 1988. Vanuatu – 2 males (37.6 × 29.0 mm, 37.0 × 28.4 mm) (ZRC 2012.103), Espiritu Santo, entrance of Loren Cave, coll. G. Boxshall, 13 September 2006. Tuvalu Islands – 1 male (42.9 × 34.3 mm) (UF 6316), among breadfruit leaves, area with coconuts, strand vegetation, 2 m asl, Vaiaku, Fongafale, Tuvalu, Funafuti Atoll, coll. L. Kirkendale, 23 June 2004. Wallis & Futuna Islands – 1 female (38.3 × 29.5 mm) (MNHN-IU-2013-13174), coll. J. Poupin, 2007. Niue – 1 male (51.7 × 41.6 mm) (UF 2192), supratidal to well inland, Hakupu area, coll. B. Holthuis and G. Paulay, 9 April 1991. Australian Seas – 1 female (holotype of Discoplax pagenstecheri Kossmann, 1878) (55.1 × 44.1 mm) (SMF 6872) (ex Mus. Heidelberg Cr190, Mus. Godeffroy). Cocos-Keeling Islands (Australia) – 1 male (46.5 × 57.4 mm) (ZRC 1965.12.1.7), coll. C.A. Gibson–Hill, 1941; 1 male, 1 female (ZRC 1965.12.1.8–9), coll. M.W.F. Tweedie, 1941; 1 male (57.6 × 44.5 mm) (ZRC 2013.1074), Horsburgh Island, coll. P.K.L. Ng et al., 23 March 2011; 6 males, 1 female (ZRC 2013.1075), Horsburgh Island, coll. P.K.L. Ng et al., 23 March 2011; 1 male (ZRC 2013.1084), station CK24, west southwest of Rumah Baru (boat launch point), West Island, coll. P.K.L. Ng, 24 March 2011; 3 males (ZRC 2019.744), within 500 m of landing point on island, North Keeling Island, coll. P.K.L. Ng, 8 February 2012; 1 female (ZRC 2019.741), within 500 m of landing point on island, North Keeling Island, coll. P.K.L. Ng, 8 February 2012; 1 female (ZRC 2019.743), station CK 2-14, National Park Camp, central west side of lagoon, North Keeling Island, coll. P.K.L. Ng, 8 February 2012; 1 male, 1 female (ZRC 2021.866) National Park Camp, central west side of lagoon, North Keeling Island, coll. P.K.L. Ng, 8 February 2012. Mauritius – 1 male (no other data), Mauritius Institute. © 2023 Academia Sinica, Taiwan © 2023 Academia Sinica, Taiwan Colour: The adult coloration of T. rotundum is surprisingly variable. Most specimens tend to have a dark brown carapace, the ambulatory legs are usually dark brown with the joints and dactylus orange, and the chelipeds are reddish-brown to purplish with palecoloured fingers (Fig. 1A, B, E, F). Some specimens, however, are much lighter in colour with yellow to orange chelipeds and ambulatory legs (Fig. 1B, C). Distribution: East Africa (Kenya), Aldabra, Seychelles, Mauritius, Cocos-Keeling Islands, Christmas Island, Japan (Ryukyus), Taiwan, Philippines, Indonesia (Ambon), Papua New Guinea, Guam, Ujae Atoll, Solomon Islands, Vanuatu, New Caledonia, Tuvalu Islands, Wallis & Futuna, Tokelau, Niue, Cook Islands. © 2023 Academia Sinica, Taiwan Remarks: This well-known species has a wide distribution in the western Indian Ocean to western Pacific (Fig. 13) and the taxonomy has been treated at length by Türkay (1974a: 234). The type of the species is still extant, with Guinot et al. (2018: 551) listing a syntype male measuring 63.0 × 50.0 mm in the MNHN as obtained from “Oceania”. Quoy and Gaimard (1824: 527) had stated in the original paper that the material was from Guam. NK Ng et al. (2019: 101) designated this dried male specimen in MNHN as the lectotype of the species (MNHN-IU-2000-3745). Our material from Guam agrees very well with their description and figure (Guinot et al. 2018: fig. 4G, NK Ng et al. 2019: 101, fig. 3B). Henri Milne Edwards (1853) described Cardisoma frontalis from an unknown locality and his description is too brief to be helpful. Most authors have treated C. frontalis as a junior synonym of C. rotundum (see Türkay 1974a b; PKL Ng et al. 2008; Guinot et al. 2018; NK Ng et al. 2019). Türkay (1974b) discussed the issue with the types of these two species, selecting the syntype of Thelphusa rotunda as the holotype, but incorrectly selecting this specimen also as the holotype of Cardisoma frontalis H. Milne Edwards, 1853. As discussed by NK Ng et al. (2019), the type of C. frontalis is lost and there is no evidence that the specimen(s) of Quoy and Gaimard (1824) and H. Milne Edwards (1853) are the same. They noted that as the synonymy of this species was well accepted and there was no problem with their taxonomy, there was no immediate need to select a neotype for Cardisoma frontalis. With the recognition of a new species from French Polynesia, there is now a problem as it is possible H. Milne Edwards’ (1853) original specimen(s) originated from there. Henri Milne Edwards’ (1853: 204) account for the species is relatively brief and not informative: “Carapace assez semblable à celle du C. carnifex par sa forme Générale et ses crêtes marginales latéro-antérieures, mais s’en distinguant, ainsi que de celle des autres espèces dont il vient d’être question, par la saillie considérable des lobes prégastriques et la profondeur du sillon mésogastrique. Basicérite étroit. Bras du mâle gros et presque de même grandeur des deux côtés; mains renflées; pinces courtes. Pattes ne portant que des poils très courts; le propodite garni en dessus d’une double rangée de petites épines. Patrie inconnue.” [Carapace quite similar to that of C. carnifex in its general shape and its anterolateral ridges, but differing from it, as well as from that of the other species just mentioned, by the considerable projection of the pregastric lobes and the depth of the mesogastric groove. Narrow baserite. Male cheliped large and nearly equal on both sides; swollen palm; short fingers. Legs with only very short hairs; dorsal margin of propodus with a double row of small spines. Unknown provenance.]. It is possible to keep the name Cardisoma frontalis H. Milne Edwards, 1853 by selecting the holotype of the new species from French Polynesia as its neotype. We have, however, decided otherwise because Cardisoma frontalis has long been confused with it and is now generally regarded as the junior synonym of Tuerkayana rotundum. In our opinion, to establish a new name would be less confusing. Kossmann (1878) described Discoplax pagenstecheri from a female specimen from Australian seas (Figs. 2F, 5F); as treated by Türkay (1974a), it is clearly identical to T. rotundum. Innocenti and Vannini (2007) reported a male specimen of T. rotundum from Mombasa, Kenya. The late Michael Türkay passed the first author photographs (Fig. 2G, H) of a specimen he had seen in the Mauritius Institute in the 1970s but has never been published. The adult male specimen (no measurements indicated) is clearly T. rotundum as defined here. Unfortunately, we were unable to obtain this specimen for study, but the photograph leaves no doubt as to its identity. The only record from Christmas Island in the Indian Ocean is a photograph of a specimen observed at the Blowholes on the island (Fig. 1F) (Orchard 2012: 149). It is surprising that it is not common on the island; it is relatively abundant on the nearby Cocos-Keeling Islands (Tweedie 1950; present data). Rathbun (1906: 838) listed two specimens of “ Cardisoma rotunda ” from Oahu in Hawai’i but there have not been any records since. Castro (2011: 113) discussed this matter at length and concluded that the original specimens had been mislabelled., Published as part of Ng and Hsi-Te Shih, Peter K. L., 2023, Tuerkayana latens, a New Species of Land Crab from French Polynesia, with a Discussion on the Phylogeny of the Genus (Crustacea: Decapoda: Brachyura: Gecarcinidae), pp. 1-21 in Zoological Studies 62 (10) on pages 2-7, DOI: 10.6620/ZS.2023.62-10, http://zenodo.org/record/8075407, {"references":["Quoy J, Gaimard JP. 1824. Zoologie. In: Freycinet LD de (ed.), Voyage autour du monde, enterpris par ordre du Roi, sours le Ministere et confromement aux instructions de S. Exc. M. le Vicomte du Bouchage, Secretaire d'etat au Department de la Marine, Execute sur les corvettes de S. M. [= Sa Majeste] l'Uranie et la Physicienne, pendant les annees 1817, 1818, 1819 et 1820. Pillet Aine, Paris. Vol 3 (1): [3] [= Preface], 1 - 376; Vol 3 (2): 377 - 712; Atlas: pls. 1 - 96. [Genre Crabe: Pp. 522 - 544, pls. 76 - 82]","Ng NK, Rodriguez Moreno PA, Naruse T, Guinot D, Mollaret N. 2019. Annotated catalogue of brachyuran types (Crustacea, Decapoda, Brachyura) in the Museum national d'Histoire naturelle, Paris. Part II. Thoracotremata, superfamily Grapsoidea (Gecarcinidae and Grapsidae). Zoosystema 41: 91 - 130. doi: 10.5252 / zoosystema 2019 v 41 a 7.","Man JG De. 1902. Die von Herrn Professor Kukenthal in Indischen Archipel gesammelten Dekapoden und Stomatopoden. Abh Senckenb Naturforsch Ges 25: 467 - 929, pls. 19 - 27.","Tweedie MWF. 1950. The fauna of the Cocos-Keeling Islands, Brachyura and Stomatopoda. Bull Raffles Mus 22: 105 - 148.","Kossmann R. 1878. Kurze Notizen uber einige neue Crustaceen sowie uber neue Fundorte einiger bereits beschriebenen. Arch Naturg 44: 251 - 258. doi: 10.5962 / bhl. part. 12966.","Turkay M. 1974 a. Die Gecarcinidae Asiens und Ozeaniens (Crustacea: Decapoda). Senck biol 55: 223 - 259.","Sakai T. 1976. Crabs of Japan and the Adjacent Seas. Kodansha Ltd., Tokyo, 773 pp., 251 pls.","Takeda M. 1982. Keys to the Japanese and Foreign Crustaceans Fully Illustrated in Colors. Hokuryukan Ltd., Tokyo, Japan, 58 + 284 pp. (in Japanese)","Miyake S. 1983. Japanese Crustacean Decapods and Stomatopods in Color. Vol. II: Brachyura (Crabs). Hoikusha, Osaka, Japan, 8 + 277 pp., 64 pls. (in Japanese)","Hirata Y, Nakasone Y, Shokita S. 1988. Okinawan Decapods and Shells in Color. Fudokisha, Okinawa, Japan, 144 pp. (in Japanese)","Ho PH, Shy JY, Yu HP. 1992. The land crabs of the family Gecarcinidae (Crustacean, Decapoda, Brachyura) of Taiwan. Ann Taiwan Mus 35: 51 - 66. (in Chinese)","Poupin J. 1994. Quelques Crustaces Decapodes communs de French Polynesia. Rapport Scientifique du Service Mixte de Surveillance Radiologique et Biologique de l'homme et de l'environnement. SMSRB, Montlhery, France, 86 pp., 68 figs., 8 pls.","Ng PKL. 1998. Crabs. In: Carpenter KE, Niem VH. (eds), FAO Species Identification Guide for Fishery Purposes. The Living Marine Resources of the Western Central Pacific. Volume 2. Cephalopods, Crustaceans, Holothurians and Sharks. Food and Agriculture Organisation (FAO), Rome, Italy, pp. 1045 - 1155.","Liu HC. 1999. Tales of Taiwanese land crabs. Taiwan Nat Sci 63: 76 - 90. (in Chinese)","Lee JH. 2001. A Field Guide to Crabs in Taiwan. Bigtrees Co., Taipei, 174 pp. (in Chinese)","Davie PJF. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda - Anomura, Brachyura. In: Wells A, Houston WWK (eds), Zoological Catalogue of Australia. CSIRO Publishing, Melbourne, Australia, 19.3 B: 641 pp.","Lin DR, Chao RF, Lu CY, Hsieh LC. 2011. Guide to the Land Crabs of Lyudao (Green Island), Taiwan. Taitung County Government, Taitung, Taiwan, 63 pp. (in Chinese)","Ng PKL, Shih HT, Ho PH, Wang CH. 2017. An updated annotated checklist of brachyuran crabs from Taiwan (Crustacea: Decapoda). J Nat Taiwan Mus 70 (3 / 4): 1 - 208. doi: 10.6532 / JNTM. 201712 _ 70 (3; 4). 01.","Innocenti G, Vannini M. 2007. The first record of the land crab, Discoplax rotunda (Quoy & Gaymard, 1824) (Decapoda, Brachyura, Gecarcinidae) from East Africa: The problem of the island species. Crustaceana 80: 635 - 637. doi: 10.1163 / 156854007780765632.","Ng PKL, Guinot D, Davie PJF. 2008. Systema Brachyurorum: Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bull Zool Suppl 17: 1 - 296.","Poupin J, Juncker M. 2008. Crustaces des iles Wallis & Futuna: inventaire illustre, especes commercialisables et capture des formes larvaires. Rapport technique du CRISP. Coral Reef Initiatives for the Pacific, Noumea, Nouvelle-Caledonie, 44 pp., 18 figs., 7 pls.","Poupin J, Juncker M. 2010. A Guide to the Decapod Crustaceans of the South Pacific. Secretariat of the Pacific Community, Noumea, New Caledonia, 317 pp.","Orchard M. 2012. Crabs of Christmas Island. Christmas Island Natural History Association, Christmas Island, Australia, 288 pp.","Li JJ, Chiu YW. 2013. Land Crabs of Hengchun Peninsula. National Museum of Marine Biology & Aquarium, Pingtung, Taiwan, 91 pp. (in Chinese)","Shih HT, Komai T, Liu MY. 2013. A new species of fiddler crab from the Ogasawara (Bonin) Islands, Japan, separated from the widely-distributed sister species Uca (Paraleptuca) crassipes (White, 1847) (Crustacea: Decapoda: Brachyura: Ocypodidae). Zootaxa 3746: 175 - 193. doi: 10.11646 / zootaxa. 3746.1.8.","Ng PKL, Shih HT. 2014. The systematics of the land crabs of the Discoplax hirtipes (Dana, 1851) species-group (Crustacea: Decapoda: Brachyura: Gecarcinidae), with description of a new species from the eastern Indian Ocean. Raffles Bull Zool Suppl 30: 109 - 135.","Toyota K, Seki S, Komai T. 2014. Freshwater Shrimps and Crabs of Japan: 102 Species from Brackish and Fresh Water. Seibundo Shinkosha, Tokyo, 256 pp. (in Japanese)","Shih HT, Saher NU, Kamrani E, Ng PKL, Lai YC, Liu MY. 2015. Population genetics of the fiddler crab Uca sindensis (Alcock, 1900) (Crustacea: Brachyura: Ocypodidae) from the Arabian Sea. Zool Stud 54: 1. doi: 10.1186 / s 40555 - 014 - 0078 - 3.","Li JJ, Chiu YW. 2019 a. An Atlas of Land Crabs on Hengchun Peninsula 2.0. National Museum of Marine Biology & Aquarium, Pingtung, Taiwan, 136 pp. (in Chinese)","Li JJ, Chiu YW. 2019 b. A Field Guide of Land Crabs on Hengchun Peninsula 2.0. National Museum of Marine Biology & Aquarium, Pingtung, Taiwan, 64 pp. (in Chinese)","Guinot D, Ng NK, Rodriguez Moreno PA. 2018. Review of grapsoid families for the establishment of a new family for Leptograpsodes Montgomery, 1931, and a new genus of Gecarcinidae H. Milne Edwards, 1837 (Crustacea, Decapoda, Brachyura, Grapsoidea MacLeay, 1838). Zoosystema 40: 547 - 604. doi: 10.5252 / zoosystema 2018 v 40 a 26.","Sasaki J. 2019. The Species List of Decapoda, Euphausiacea, and Stomatopoda, all of the World, version 03 - 3.1. Local Independent Administrative Agency Hokkaido Research Organization, Resources Management and Enhancement Division, Abashiri Fisheries Research Institute, Fisheries Research Department, Hokkaido, Japan, 14644 pp. doi: 10.13140 / RG. 2.2.22353.89446.","Toyota K, Seki S, Komai T. 2019. Fresh and Brackish Water Shrimps and Crabs of Japan. Midori-Shobo, Tokyo, 339 pp. (in Japanese)","Milne Edwards H. 1853. Memoire sur la famille des Ocypodides. Suite (1). Deuxieme Tribu Principale. Ann Sci Nat Zool 20: 163 - 228, pls. 6 - 11.","Turkay M. 1974 b. Bemerkungen zu einigen Landkrabben (Crustacea, Decapoda). Bull Mus natn Hist nat Paris (3) 142: 969 - 980 (dated 1973).","Rathbun MJ. 1906. The Brachyura and Macrura of the Hawaiian islands. Bull US Fish Comm 23: 827 - 930, pls. 1 - 24.","Castro P. 2011. Catalog of the anomuran and brachyuran crabs (Crustacea: Decapoda: Anomura, Brachyura) of the Hawaiian Islands. Zootaxa 2947: 1 - 154. doi: 10.11646 / zootaxa. 2947.1.1."]}

Austruca perplexa (H. Milne Edwards, 1852) Fig. 5G Uca (Celuca) lactaea perplexa, Crane 1975: 300, 613, figs. 18D–F, 19C, 26D, 31E, 54K, KK, 69C.— Manuel et al. 1991: 84. Material examined. 2 males, 15.7 by 9.5 mm, 16.3 by 10.1 mm (ZRC), Bgy. Baluarte, Tagoloan town, Misamis Oriental prov., coll. N.J. Lagare, 23 Sep. 2016. Remarks. This species was described as Gelasimus perplexus by H. Milne Edwards (1852) from Java, Indonesia. In her revision of the fiddler crabs (Uca) of the world, Crane (1975) treated it as a subspecies of Uca (Celuca) lactaea, and recorded it from several localities in the Philippines, including Mindanao (see Table 1). Subsequent workers (e.g. Ng et al. 2008; Shih et al. 2009; Naderloo et al. 2010) treated it as a full species, and when Shih et al. (2016) did their phylogenetic study of Ocypodidae, they clarified the taxonomy of the previously convoluted subgeneric classification of the genus Uca, and recognized many of the subgenera as full genera. The so-called “ Uca lactaea ” species complex (which included “ Uca ” perplexa) was shown to be part of a monophyletic clade and was included in the genus Austruca Bott, 1973 (whose composition was nearly identical to Crane’s (1975) concept of Indo-West Pacific Celuca) (Shih et al. 2016). The present material was collected from the estuary of the Tagoloan River, and agrees well with the published keys, descriptions and illustrations of A. perplexa (cf. Crane 1975: 300, Naderloo et al. 2010: 24, fig. 16, 18c, f; Shih et al. 2016: 62, fig. 2C–E)., Published as part of Lagare, Nickel Jean S., Mapi-Ot, Emmarie F., Molina, Ziljih S., Neri, Jemateo B., Nuñeza, Olga M. & Mendoza, Jose Christopher E., 2020, On a collection of freshwater and estuarine crabs (Crustacea: Brachyura) from Mindanao Island, the Philippines, pp. 301-330 in Zootaxa 4868 (3) on page 319, DOI: 10.11646/zootaxa.4868.3.1, http://zenodo.org/record/4443564, {"references":["Crane, J. (1975) Fiddler Crabs of the World. Ocypodidae: Genus Uca. Princeton University Press, New Jersey, xxiii + 736 pp.","Manuel, M. R., Gonzales, P. C. & Basmayor, L. (1991) The crustacean fauna of Natunawan Cove in Tabaco, Albay (Crustacea: Brachyura, Macrura and Stomatopoda). National Museum Papers, 2 (1), 59 - 102.","Shih, H. - T., Kamrani, E., Davie, P. J. F. & Liu, M. Y. (2009) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U. lactea species-complex. Hydrobiologia, 635, 373 - 382. https: // doi. org / 10.1007 / s 10750 - 009 - 9930 - 6","Naderloo, R., Turkay, M. & Chen, H. - L. (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 (1), 1 - 38. https: // doi. org / 10.11646 / zootaxa. 2500.1.1"]}

Zeylanicobdella arugamensis de Silva, 1963 General distribution and habitats. Distributed in Iranian coast of Persian Gulf and Indian Ocean (Raj 1974; Polgar et al. 2009). Usually found living as ecto-parasite on mudskippers. Remarks. Reported by Polgar et al. (2009) from Iranian coast of Persian Gulf. A summary of all known leech species from Iran is presented in Table 1. Family Genus Species Arhynchobdellids Erpobdellidae Dina D. farsa * D. lineata concolor * D. lineata lineata * D. orientalis D. punctata punctata * + D. stschegolewi * Erpobdella E. borisi * E. monostriata * + E. octoculata * E. vilnensis * + Trocheta Tr. haskonis * + Haemopidae Haemopis Ha. sanguisuga * Hirudinidae Hirudo Hi. orientalis * Limnatis L. nilotica L. paluda * Rhynchobdellids Glossiphoniidae Glossiphonia G. concolor * + Helobdella Hel. stagnalis * Hemiclepis Hem. marginata Placobdella Pl. costata * Theromyzon Th. affinis Ozobranchidae Ozobranchus O. branchiatus Piscicolidae Piscicola Pi. geometra Zeylanicobdella Z. arugamensis, Published as part of Darabi-Darestani, Kaveh, Sari, Alireza & Sarafrazi, Alimorad, 2016, Five new records and an annotated checklist of the leeches (Annelida: Hirudinida) of Iran, pp. 41-70 in Zootaxa 4170 (1) on page 67, DOI: 10.11646/zootaxa.4170.1.2, http://zenodo.org/record/266620, {"references":["Raj, P. J. (1974) A review of the fish-leeches of the Indian Ocean. Journal of the Marine Biological Association of India, 16, 381 - 397.","Polgar, G., Burreson, E. M., Stefani, F. & Kamrani, E. (2009) Leeches on Mudskippers: Host - Parasite Interaction at the Water's Edge. Journal of Parasitology, 95 (4), 1021 - 1025. http: // dx. doi. org / 10.1645 / GE- 1718.1"]}

Uca (Petruca) panamensis (Stimpson, 1859) (Figs. 1 ���5, 8) Gelasimus panamensis Stimpson, 1859: 63; Smith 1870: 139, pl. 4 (5); Kingsley 1880: 150, fig. 10 (24); Cano 1889: 235. Uca panamensis ��� Nobili 1901: 49; Rathbun 1918: 412, pl. 149; Crane 1941: 204, test-fig. 4 X; Garth 1948: 60; Takeda & Murai 2003: 179, fig. 1. Uca (Minuca) panamensis ��� Bott 1954: 162, fig. 4, pl. 15 (4); Crane 1975: 158, pl. 22 A���D, figs. 46 H, 66 G; Prahl & Guhl 1979: 147, figs. 1���2. Uca (Leptuca) panamensis ��� Rosenberg 2001: 848, 852; Beinlich & von Hagen 2006: 22, 26, fig. 7 j; Ng et al. 2008: 241. Material examined. Neotype for Gelasimus panamensis, 1 ♂ (16.8 mm), USNM 1294205 (transferred from NCHUZOOL 13581), Culebra Island, Panam��, coll. J. Christy, 2008; 1 ♂ (13.0 mm), NCHUZOOL 13581, same data as neotype; 1 ♂ (12.6 mm), ZRC 2012.0126, Playa San Juanillo, Costa Rica, coll. Z. Jaafar, 25���27 July 2011; 4 ♂♂ (11.1���15.1 mm), 2 ♀♀ (11.7, 13.7 mm), ZRC, 1 ♂ (16.4 mm), 1 ♀ (16.8 mm), NCHUZOOL 14753, San Juanillo rocky shore, Ostional, Costa Rica, coll. Z. Jaafar, 26���29 July, 2011. Comparative material. See Appendix 1. Remarks. Gelasimus panamensis was described by Stimpson (1859) from three specimens, all apparently immature, from an undisclosed site in Panam��. No figures were provided. In any case, the types are no longer extant. Like most of his material, they were almost certainly lost in the great Chicago Fire (see Evans 1967; Deiss & Manning 1981; Manning 1993; Vasile et al. 2005; Manning & Reed 2006). Although the species has been well treated by many authors, notably Crane (1975), since a new subgenus is established here for the species, the designation of a neotype for Gelasimus panamensis Stimpson, 1859, is important for longer term taxonomic stability. This is especially important given that there is now active research on Uca. To this effect, we here select an adult male with CW of 16.8 mm collected from Panam�� as the neotype of the species. It is a recently collected specimen (USNM 1294205) that also has genetic data. DNA analyses and discussion. A 589 bp segment of the 16 S, 658 bp segment of COI and 689 bp segment of 28 S, from 55 species of fiddler crabs, were amplified and aligned (Table 1). The phylogenetic tree of the combined markers was reconstructed from the BI analysis, with the support values from two methods of ML analyses (Fig. 9). To estimate the divergence time, the substitution rate of 1.17 % / 10 6 year for the combined 16 S and COI of marine sesarmids (cf. Schubart et al. 1998; Ragionieri et al. 2009) was applied to the p-distance between clades. The relationships of the IWP subgenera (Fig. 9) generally agree with Shih et al. (2013 b) and Shih (2015) and are not repeated here. The American broad-fronted species form a highly-supported main clade by three methods, with three distinct clades corresponding to three subgenera with high support at least by the BI method. The subgenus Petruca subgen. nov. diverged earlier than the others at about 14 million years ago (mya) (pdistance of the combined 16 S and COI = 16.45 %). Both Minuca and Leptuca are monophyletic, are closely related and diverged at 9 mya (p-distance = 10.63 %). The six specimens of U. panamensis from the Pacific coasts of Costa Rica and Panam�� form a highly supported clade supporting the establishment of Petruca subgen. nov. Its intraspecific K 2 P divergence COI is average 1.73 % (0.33���2.81 %), which is lower than U. splendida (1.86 %; cf. Shih et al. 2012), but higher than other reported fiddler crabs. For example, the largest intraspecific divergences are 1.23 % in U. jocelynae (recalculated from Shih et al. 2010); 0.98 % in U. annulipes (Shih et al. 2009); about 0.83 % in U. pugnax (cf. Sanford et al. 2006); 0.46 % in U. sindensis (recalculated from Shih et al. 2015); about 0.32 % in U. maracoani (cf. Wieman et al. 2014); and about 0.13 % in U. annulipes (= U. aff. annulipes, see Shih et al. 2013 b) from East Africa (Silva et al. 2010). The high genetic variability within a species may be explained by past historical or present barriers (e.g., ocean currents and salinity) affecting their larval dispersal (see L��pez-Duarte et al. 2011; Shih et al. 2015). Within the Minuca clade, U. brevifrons diverged earliest at 8 mya (p-distance = 9.27 %), and is sister to the remaining species of this subgenus. The habitat of U. brevifrons is unusual. This species lives on the muddy banks of freshwater and brackish streams, sometimes even in wet forest several meters from the nearest stream (Crane 1941, 1975). Perhaps the simpler form of urocardiac ossicles of gastric mill present in this species (Fig. 6 A) is an ancestral character. Crane (1975) treated U. brevifrons and U. minax as the same ���superspecies [minax]��� but their close relationship is not supported genetically. The taxonomy of Uca virens Salmon & Atsaides, 1968, and U. longisignalis Salmon & Atsaides, 1968, has been debated since the publication of the description of the two species (Salmon & Atsaides 1968). By comparing the holotypes of the above two species, von Hagen (1980) confirmed them as synonyms of U. rapax and U. minax, respectively. The synonymization of U. virens as U. rapax has been followed by later studies (e.g. Barnwell & Thurman 1984; Beinlich & von Hagen 2006). Although there are several studies still using the name of ��� Uca virens ��� (Rosenberg 2001; Swanson et al. 2013; P��rez-Mozqueda et al. 2014; del Castillo et al. 2015), the identification of the species in these studies will need to be reconfirmed (C. Thurman, pers. comm.). Based on the allozyme study of hemolymph amylases (Barnwell & Thurman 1984) and esterase- 1 and - 2 (Salmon & Kettler 1987), it is apparent that there are two ���forms��� of U. rapax, corresponding to the distribution of U. virens and U. rapax. In our study based on 16 S, COI and 28 S markers (Fig. 9), there is a clade sister to U. rapax with similar morphology. Before additional studies are done to confirm whether it is actually U. virens, we use the name ��� U. cf. virens ��� for convenience. The two species diverged at 2 mya (p-distance = 2.32 %), although Salmon & Kettler (1987) estimated the divergence time is 20���50 thousand years based on the allozyme data. While previous studies considered the morphological characters of U. virens were within the range of U. rapax (Hagen 1980; Barnwell & Thurman 1984), their gastric mill structures show differences in the posterior tooth plate of urocardiac ossicles, with 8 ridges in U. rapax (Fig. 6 F) and 6 ridges in U. virens (Fig. 6 B). More studies of morphological characters, especially the gastric mills, are necessary to clarify the identity of U. virens. Another confused species is U. longisignalis, which has been synonymized as U. minax by von Hagen (1980). Thurman (1982) considered it is a valid species because there are substantial morphological differences between the two species, a decision that has been followed by other studies thereafter. Genetically, Felder & Saton (1994) showed that the allozyme evidence did not support the hypothesis that U. longisignalis is different from U. minax. Landstorfer et al. (2008) and Warwick et al. (2009) also reported the failure to separate the two species by mitochondrial COI or nuclear internal transcribed spacer (ITS- 1) markers. We included in our study specimens of the two species with typical characters of each taxon (mainly based on with or without pubescence ventrally; cf. Barnwell & Thurman 1984), but the molecular data only shows one clade (Fig. 9), therefore agreeing with earlier genetic studies. Further studies, with more complete sampling, will be necessary to clarify the taxonomy of the U. minax complex. The close relationship of U. burgersi and U. mordax suggested by Salmon & Kettler (1987) is highly supported (Fig. 9). Within the Leptuca clade, U. panacea, and U. pugilator are closely related, with the divergence time at 2.8 mya (p-distance = 3.24 %), which supports the original hypothesis that both species have diverged recently (Novak & Salmon 1974; Barnwell & Thurman 1984). The supposed close relationship of U. speciosa and U. spinicarpa (Crane 1975; Barnwell & Thurman 1984) is not supported in this study (Fig. 9). Our study, however, agrees well with the allozyme studies (Salmon et al. 1979; Salmon & Kettler 1987) that the genetic distance of two species is large. Uca speciosa is distinct from other species genetically, although it forms a main clade, with U. dorotheae, U. deichmanni, and U. cumulanta. Uca spinicarpa is, however, closely related to the species pair, U. thayeri and U. umbratila (see below). Crane���s (1975) subgenus Boboruca (= Planuca Bott, 1973), established for U. thayeri and U. umbratila, was synonymized with Minuca by Albrecht & von Hagen (1981), based on combined allozyme and morphological characters, and this has been followed by all later authors (e.g., von Hagen 1987; Salmon & Zucker 1988; Rosenberg 2001; Beinlich & von Hagen 2006; Ng et al. 2008). In this study, U. thayeri and U. umbratila came out clearly inside the Leptuca clade (Fig. 9). In the studies of Levinton et al. (1996) and Sturmbauer et al. (1996) using the 16 S marker, the position of U. umbratila is within a ���derived American clade��� composed of a mixture of the subgenera Minuca and Leptuca. Uca umbratila is nevertheless located within Minuca in Landstorfer & Schubart's (2010) study, although their sequences of COI (FN 430708) and 28 S (FN 430719) agree well with U. umbratila in our study. The different conclusions are probably caused by the inclusion of 16 S marker, as well as the number of species used in ingroups and outgroups. Hampton et al. (2014) have pointed out the unsolved relationship of Boboruca from the above studies. Our study gives genetic support for including U. thayeri and U. umbratila in the subgenus Leptuca (Fig. 9), although their morphology is closer to Minuca. One of the main characters placing these two species in Minuca is the presence of two posterolateral striae on each side of carapace (cf. Beinlich & von Hagen 2006; Bezerra 2012). This character, however, can be variable. While there are two striae on each side of the carapace in U. thayeri, U. umbratila only has one stria (cf. Crane 1941: text-fig. 7). This has not been discussed before. The available data also implies the morphological similarity between the two species and Minuca is convergent. The present study shows that U. thayeri, U. umbratila, and U. spinicarpa form a highly supported clade which may be the result of allopatric speciation. Whereas U. thayeri and U. umbratila diverged at 3.7 mya (p-distance = 4.34 %), a well-known species pair separated after the closure of the Isthmus of Panam�� around 3.1 million years ago (reviewed by Schubart et al. 1998; Lessios 2008), U. spinicarpa is endemic to the Gulf of Mexico in a more northern distribution, and it diverged from the other two species at 6.3 mya (p-distance = 7.38 %). Uca subcylindrica is endemic along the border between Texas and northern Mexico in the Gulf of Mexico, and has been considered as an anomalous form, with unusually rounded body, striking form of G 1 and hypersaline lagoonal habitat, without close alliance to other species (Crane 1975; Barnwell & Thurman 1984; Thurman 1987). Crane (1975) placed it in Minuca, but noted with reservation that it was close to Leptuca. Because some characters of U. subcylindrica are shared with several members of Leptuca, Barnwell & Thurman (1984) considered it was not suitable to retain in the homogeneous Minuca and transferred it to Leptuca. Their action is supported by our study, as U. subcylindrica forms a distinct clade from other species of this subgenus (Fig. 9). The simpler form of urocardiac ossicles of the gastric mill (Fig. 7 A) may also be a reflection of the early split of this species from others. Uca panamensis was placed in Minuca mainly by the characters of the front of the carapace and male first gonopod (Bott 1954, 1973; Crane 1975). Based on a morphological analysis, Rosenberg (2001) placed it in Leptuca along with U. terpsichores, U. deichmanni, U. leptodactyla, and U. dorotheae. The many unusual features of U. panamensis (e.g., flatter carapace, stiff setal brushes on the tips of the small chelae, the posterior extension of major manus, preference for rocky habitats etc.) suggest this classification is incorrect. The present use of new morphological characters (viz. the orbital armature at the inner corner of orbital floor, simple form of urocardiac ossicles of gastric mill, large and bent inner process of the G 1) as well as molecular evidence using mitochondrial and nuclear markers (Fig. 9), support the hypothesis that U. panamensis belongs to its own subgenus (Petruca subgen. nov.), and diverged from other American Minuca and Leptuca at about 14 mya., Published as part of Shih, Hsi-Te, Ng, Peter K. L. & Christy, John H., 2015, Uca (Petruca), a new subgenus for the rock fiddler crab Uca panamensis (Stimpson, 1859) from Central America, with comments on some species of the American broad-fronted subgenera, pp. 471-494 in Zootaxa 4034 (3) on pages 486-489, DOI: 10.11646/zootaxa.4034.3.3, http://zenodo.org/record/235573, {"references":["Stimpson, W. (1859) Notes on North American Crustacea, No. 1. Annals of the Lyceum of Natural History of New York, 7 (4 - 5), 49 - 93, pl. I.","Smith, S. I. (1870) Notes on American Crustacea. No. 1. Ocypodoidea. Transactions of the Connecticut Academy of Arts and Science, 2, 113 - 176, pls. 2 - 5.","Kingsley, J. S. (1880) Carcinological notes, no. II. - Revision of the Gelasimi. Proceedings of the Academy of Natural Sciences of Philadelphia, 1880, 135 - 155.","Cano, G. (1889) Crostacei brachiuri ed anomuri raccolti nel viaggio della \" Vettor Pisani \" intorno al globo. Bollettino della Societa di Naturalisti in Napoli, 3, 79 - 106, 169 - 268.","Nobili, G. (1901) Viaggio del Dr. Enrico Festa nella Republica dell'Ecuador e regione vicine. Decapodi e Stomatopodi. Bollettino dei Musei di Zoologia ed Anatomia comparata della R. Universit di Torino, 16 (415), 1 - 58.","Rathbun, M. J. (1918) The grapsoid crabs of America. Bulletin of the United States National Museum, 97, 1 - 461, pls. 1 - 161. http: // dx. doi. org / 10.5479 / si. 03629236.97. i","Crane, J. (1941) Crabs of the genus Uca from the west coast of Central America. Eastern Pacific Expeditions of the New York Zoological Society. XXVI. Zoologica, 26 (19), 145 - 208.","Garth, J. S. (1948) The Brachyura of the \" Askoy \" Expedition, with remarks on carcinological collecting in the Panama Bight. Bulletin of the American Museum of Natural History, 92, 1 - 66.","Takeda, S. & Murai, M. (2003) Morphological and behavioural adaptations to the rocky substrate by the fiddler crab Uca panamensis (Stimpson, 1859): preference for feeding substratum and feeding mechanism. Journal of Experimental Marine Biology and Ecology, 287, 179 - 191. http: // dx. doi. org / 10.1016 / S 0022 - 0981 (02) 00550 - 6","Bott, R. (1954) Dekapoden (Crustacea) aus El Salvador. 1. Winkerkrabben (Uca). Senckenbergiana Biologica, 35 (3 / 4), 155 - 180.","Crane, J. (1975) Fiddler Crabs of the World (Ocypodidae: Genus Uca). Princeton University Press, Princeton, New Jersey, xxiii + 736 pp.","Prahl, H. von & Guhl, F. (1979) Notas sobre la biologia de Uca (Minuca) panamensis (Stimpson, 1859) (Crustacea, Ocipodidae) y su distribucion en Colombia. Anales del Instituto de Investigaciones Marinas de Punta Betin, 11, 147 - 152.","Rosenberg, M. S. (2001) The systematics and taxonomy of fiddler crabs: a phylogeny of the genus Uca. Journal of Crustacean Biology, 21, 839 - 869. http: // dx. doi. org / 10.1163 / 20021975 - 99990176","Beinlich, B. & von Hagen, H. O. (2006) Materials for a more stable subdivision of the genus Uca Leach. Zoologische Mededelingen, 80, 9 - 32.","Evans, A. C. (1967) Syntypes of Decapoda described by William Stimpson and James Dana in the collections of the British Museum (Natural History). Journal of Natural History, 1, 399 - 411. http: // dx. doi. org / 10.1080 / 00222936700770391","Deiss, W. A. & Manning, R. B. (1981) The fate of the invertebrate collections of the North Pacific Exploring Expedition, 1853 - 1856. In: Wheeler, A. & Price, J. H. (Eds), History in the Service of Systematics. Society for the Bibliography of Natural History, London, pp. 79 - 85.","Manning, R. B. (1993) The scientific contributions of William Stimpson, an early American naturalist and taxonomist. Crustacean Issues, 8, 109 - 117.","Vasile, R. S., Manning, R. B. & Lemaitre, R. (2005) William Stimpson's Journal from the North Pacific Exploring Expedition, 1853 - 1856. Crustacean Research Special, 5, 1 - 220.","Manning, R. B. & Reed, K. J. (2006) Decapod crustaceans deposited in the Zoological Museum of Copenhagen by William Stimpson in 1859. Raffles Bulletin of Zoology, 54, 283 - 293.","Schubart, C. D., Diesel, R. & Hedges, S. B. (1998) Rapid evolution to terrestrial life in Jamaican crabs. Nature, 393, 363 - 365. http: // dx. doi. org / 10.1038 / 30724","Ragionieri, L., Fratini, S, Vannini, M. & Schubart, C. D. (2009) Phylogenetic and morphometric differentiation reveal geographic radiation and pseudo-cryptic speciation in a mangrove crab from the Indo-West Pacific. Molecular Phylogenetics and Evolution, 52, 825 - 834.","Shih, H. - T., Ng, P. K. L. & Liu, M. - Y. (2013 b) Systematics of the Indo-West Pacific broad-fronted fiddler crabs (Crustacea: Ocypodidae: genus Uca). Raffles Bulletin of Zoology, 61, 641 - 649.","Shih, H. - T., Naruse, T. & Ng, P. K. L. (2010) Uca jocelynae sp. nov., a new species of fiddler crab (Crustacea: Brachyura: Ocypodidae) from the Western Pacific. Zootaxa, 2337, 47 - 62.","Shih, H. - T., Kamrani, E., Davie, P. J. F. & Liu, M. - Y. (2009) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U.","Sanford, E., Holzman, S. B., Haney, R. A., Rand, D. M. & Bertness, M. D. (2006) Larval tolerance, gene flow, and the northern geographic range limit of fiddler crabs. Ecology, 87, 2882 - 2894. http: // dx. doi. org / 10.1890 / 0012 - 9658 (2006) 87 [2882: LTGFAT] 2.0. CO; 2","Wieman, A C., Berendzen, P. B., Hampton, K. R., Jang, J., Hopkins, M. J., Jurgenson, J., McNamara, J. C. & Thurman, C. L. (2014) A panmictic fiddler crab from the coast of Brazil? Impact of divergent ocean currents and larval dispersal potential on genetic and morphological variation in Uca maracoani. Marine Biology, 161, 173 - 185.","Silva, I. C., Mesquita, N. & Paula, J. (2010) Lack of population structure in the fiddler crab Uca annulipes along an East African latitudinal gradient: genetic and morphometric evidence. Marine Biology, 157, 1113 - 1126.","Lopez-Duarte, P. C., Christy, J. H. & Tankersley, R. A. (2011) A behavioral mechanism for dispersal in fiddler crab larvae (genus Uca) varies with adult habitat, not phylogeny. Limnology and Oceanography, 56, 1879 - 1892. http: // dx. doi. org / 10.4319 / lo. 2011.56.5.1879","Salmon, M. & Atsaides, S. P. (1968) Behavioral, morphological and ecological evidence for two new species of fiddler crabs (genus Uca) from the Gulf coast of the United States. Proceedings of the Biological Society of Washington, 81, 275 - 290.","Von Hagen, H. - O. (1980) A key to the \" x-species \" of North American fiddler crabs (genus Uca). Zoologische Mededelingen, Leiden, 55 (8), 87 - 96.","Barnwell, F. H. & Thurman, C. L. (1984) Taxonomy and biogeography of the fiddler crabs (Ocypodidae: genus Uca) of the Atlantic and Gulf coasts of eastern North America. Zoological Journal of the Linnean Society, 81, 23 - 87.","Swanson, B. O., George, M. N., Anderson, S. P. & Christy, J. H. (2013) Evolutionary variation in the mechanics of fiddler crab claws. BMC Evolutionary Biology, 13, 137. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 13 - 137","Perez-Mozqueda, L. L., Del Castillo-Falconi, V. & Luis Bortolini-Rosales, J. (2014) Registros adicionales del genero Uca (Brachyura: Ocypodidae) en la laguna de Tamiahua, Veracruz, Mexico. [Additional records of the genus Uca (Brachyura: Ocypodidae) in Tamiahua Lagoon, Veracruz, Mexico.] Revista Mexicana De Biodiversidad, 85, 969 - 971. http: // dx. doi. org / 10.7550 / rmb. 36600","Salmon, M. & Kettler, M. K. (1987) The importance of behavioral and biochemical differences between fiddler crab taxa, with special reference to Uca rapax (Smith) and U. virens (Salmon and Atsaides). Contributions in Marine Science, 30, 63 - 76.","Thurman, C. L. (1982) On the distinctness of the fiddler crabs Uca minax (Leconte) and Uca longisignalis Salmon & Atsaides in their region of Sympatry (Decapoda Brachyura, Ocypodidae). Crustaceana, 43, 37 - 50.","Felder, D. L. & Staton, J. L. (1994) Genetic differentiation in trans-Floridian species complexes of Sesarma and Uca (Decapoda: Brachyura). Journal of Crustacean Biology, 14, 191 - 209. http: // dx. doi. org / 10.2307 / 1548900","Landstorfer, R. B., Schubart, C. D. & Felder, D. L. (2008) Barcoding gene COI fails to distinguish between two fiddler crabs (Brachyura: Ocypodidae: Uca) across their entire range of geographic overlap. Annual Meeting, The Crustacean Society, Galveston, Texas.","Warwick, A. R., Hopkins, M. J., Berendzen, P. B. & Thurman, C. L. (2009) Morphological, physiological, and genetic variation in the red-jointed fiddler crab, Uca minax (Le Conte). Annual Meeting, Society for Integrative and Comparative Biology, Boston, MA, USA","Novak, A. & Salmon, M. (1974) Uca panacea, a new species of fiddler crab from the Gulf coast of the United States. Proceedings of the Biological Society of Washington, 87, 313 - 326.","Salmon, M., Ferris, S. D., Johnston, D., Hyatt, G. & Whitt, G. S. (1979) Behavioral and biochemical evidence for species distinctiveness in the fiddler crabs, Uca speciosa and U. spinicarpa. Evolution, 33, 182 - 191. http: // dx. doi. org / 10.2307 / 2407375","Bott, R. (1973) Die verwandtschaftlichen Beziehungen der Uca Arten. Senckenbergiana Biologica, 54, 315 - 325.","Albrecht, H. & von Hagen, H. O. (1981) Differential weighting of electrophoretic data in crayfish and fiddler crabs (Decapoda: Astacidae and Ocypodidae). Comparative Biochemistry and Physiology, 70 B, 393 - 399. http: // dx. doi. org / 10.1016 / 0305 - 0491 (81) 90272 - 8","Von Hagen, H. - O. (1987) Morphologie und Winkbalz einer neuen Uca - Art (Crustacea, Brachyura) aus dem Staat Espirito Santo (Brasilien). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 84, 81 - 94.","Salmon, M. & Zucker, N. (1988) Interpreting differences in the reproductive behaviour of fiddler crabs (genus Uca). In: Chelazzi, G. & Vannini, M. (Eds.), Behavioral Adaptation to Intertidal Life. Plenum Press, New York, pp. 387 - 407. http: // dx. doi. org / 10.1007 / 978 - 1 - 4899 - 3737 - 7 _ 26","Bezerra, L. E. A. (2012) The fiddler crabs (Crustacea: Brachyura: Ocypodidae: genus Uca) of the South Atlantic Ocean. Nauplius, 20, 203 - 246. http: // dx. doi. org / 10.1590 / S 0104 - 64972012000200011","Lessios, H. A. (2008) The Great American Schism: Divergence of marine organisms after the rise of the Central American Isthmus. Annual Review of Ecology and Systematics, 39, 63 - 91. http: // dx. doi. org / 10.1146 / annurev. ecolsys. 38.091206.095815","Thurman, C. L. (1987) Fiddler crabs (genus Uca) of eastern Mexico (Decapoda, Brachyura, Ocypodidae). Crustaceana, 53, 94 - 105. http: // dx. doi. org / 10.1163 / 156854087 X 00664"]}

Uca (Xeruca) formosensis Rathbun, 1921 (Figs. 2���8) Uca formosensis Rathbun, 1921: 155; Maki & Tsuchiya 1923: 205, pl. 23 (4); Barnwell 1982: 79; Shih 1988: 105, fig. 9, pls. 11, 12; 1997: 68, figs. 5, 17, 41; 2000: 74, 4 unnumbered figs; Fukui et al. 1989: 227, fig. 3; Huang et al. 1989: 199, fig. 8, pl. 4 D���F; Wang & Liu 1996 a: 50, 1 unnumbered fig.; 1996 b: 73, figs. 66���70; 1998: 73, figs. 66���70; 2003: 73, figs. 66���70; Ho & Hung 1997: 54; Shih et al. 1999: 164, figs. 1���4; Hung 2000: 140, fig. 440; Lee & Tung 2000: 34, 38, 7 unnumbered figs; Chen 2001: 202, fig. 637 a, b; Lee 2001: 102, 3 unnumbered figs; Ng et al. 2001: 37; Rosenberg 2001: 848, 851, 860, 866; Shen & Jeng 2005: 164, 2 unnumbered figs; Beinlich & von Hagen 2006: 25; Wang 2009: 107 ���109, 2 unnumbered figs; Liu & Wang 2010: 37, 5 unnumbered figs; Lee et al. 2013: 68, 2 unnumbered figs. Gelasimus formosensis ��� Sakai 1939: 620, text-fig. 94 b, pl. 105 (1); Horikawa 1940: 28; Lin 1949: 26. Uca (Thalassuca) formosensis ��� Crane 1975: 83, fig. 63 C, pl. 14 A���D; Sakai 1976: 604, text-fig. 331; Su & Lue 1984: 64, fig. 5; Dai et al. 1986: 425, fig. 235, pl. 59 (1); Dai & Yang 1991: 465, fig. 235, pl. 59 (1); Shih 1994: 82, figs. 15, 16, 56���59; Ho 1996: 9, figs. 1���7. Uca (Tubuca) formosensis ���Ng et al. 2008: 242. Material examined: holotype ♂ (28.8 mm), paratype ♂ (27.6 mm), USNM 54472, Rokko (= Lugang), Changhua, Taiwan, coll. Taihoku Normal School, Aug. 1919. Others: see Appendix 1. Comparative material: see Appendix 2. DNA analyses and discussion. A 578 bp segment of the 16 S, 658 bp segment of COI and 633 bp segment of 28 S from 26 species of fiddler crabs were amplified and aligned (Table 1). The phylogenetic tree of the combined markers was reconstructed from Bayesian inference analysis, with the support values from the maximum likelihood analysis (Fig. 9). It is clear that most subgenera (including Xeruca subgen. nov.) are highly supported by both methods. Xeruca is sister to Tubuca and Australuca. While Australuca is monophyletic, Tubuca becomes parapheletic. The ��� Tubuca + Australuca ��� clade, however, has high support. Gelasimus is sister to Cranuca, at least highly supported by Bayesian inference method. The clade of Austruca is monophyletic, has a high support value by Bayesian inference, with medium support by maximum likelihood. The American subgenus Uca is closely related to Afruca from eastern Atlantic. After Crane (1975), the systematics of fiddler crabs has been revised by Rosenberg (2001) and Beinlich & von Hagen (2006), with the IWP broad-fronted groups studied by Shih et al. (2009 a, 2013 b) and Naderloo et al. (2010). Uca formosensis was established by Rathbun (1921), but only less than a dozen museum specimens could be examined until the 1980 s (Crane 1975: 83; Barnwell 1982) and Crane mentioned it as an ���elusive species.��� Shih et al. (1999) revised and updated the taxonomy and morphology of the species, as well as adding ecological information after finding its preferred habitat, and examined more than 400 specimens. They suggested this species is closer to Uca (Tubuca), instead of the original designation under Uca (Thalassuca) Crane, 1975 (= Uca (Gelasimus) Bott, 1973), but notes the need for more evidence to confirm the status. Rosenberg���s (2001) detailed morphological analyses on fiddler crabs also found that U. formosensis does not clearly belong to any of the known subgenera, although closer to Tubuca and Australuca. TABLE 1. Haplotypes of 16 S rDNA, COI and 28 S rDNA for the subgenus Xeruca subgen. nov. and other related subgenera used in this study. See Material and methods for abbreviations of institutions. Madagascar: Sarodrano ZRC THH 04- 17 AB 535405 LC053377 LC053396 Cranuca U.inversa Kenya: Gazi MZUF 1024 AB 813658 AB 813674 AB 813703 Ghana: Elmina 13654 LC053362 LC053380 LC053399 The new subgenus of Xeruca is established herein for Uca formosensis based on a suite of morphological characters, including carapace, major chela, handedness, G 1 and gastric mill, which are supported by the phylogenetic analyses using mitochondrial and nuclear markers. Previous studies on the morphology showed that U. formosensis is closer to Tubuca and Australuca (Shih et al. 1999; Rosenberg 2001), which is also revealed by the molecular evidence (Fig. 9). Xeruca can be easily separated from Tubuca by the carapace shape (Fig. 4) and the fingers of the chela (Fig. 5). Although Xeruca is more similar to Australuca, the latter can be distinguished by its smaller adult size, relatively shorter dorsolateral margins, and the possession of a longer gape between the closed fingers of the adult major chela (Figs. 4, 5). The molecular evidence (Fig. 9) shows that the subgenera Tubuca and Australuca form an unresolved clade in conflict with the present taxonomy based on morphological characters (Crane 1975; Rosenberg 2001), although the Australuca species belong to their own clade. It is likely that Australuca Crane, 1975, is a synonym of Tubuca Bott, 1973. Further studies that include more species into the analysis may clarify the problem. The morphology of gastric mills is related to different functions of feeding and seems unsuitable for phylogenetic study. This may be true for the lateral gastric teeth, but not for the foregut ossicles (Br��sing & T��rkay 2011), which has been reflected by other studies: e.g. Yang (1986) on various crabs, especially Macrophthamidae; Sakai et al. (2006) on the species of the Helice and Chasmagnathus complex; Br��sing (2010) on several brachyuran families; and Naderloo et al. (2010) on the Uca lactea species complex. Based on the systematic study of the morphology of the gastric mills of IWP Uca species (in preparation), some groups have unique characters and could be used for the systematics of fiddler crabs. As mentioned above, there are several consistent characters for each of the following subgenera: Gelasimus, Uca, and Afruca (Fig. 8). Uca formosensis was published in 1921, however, its systematic position was uncertain, either in Uca (Gelasimus) (Crane 1975) or Uca (Tubuca) (Shih et al. 1999). The morphological analyses by Rosenberg (2001) have shown this species is substantially different from other species and should be placed in its own subgenus. According to the evidence of morphology and DNA sequences of this study, as well as the peculiar ecology and reproduction behavior (Shih et al. 2005), the establishment of a new subgenus Xeruca is appropriate to solve the uncertain position of this species. The monotypic Xeruca becomes a subgenus with the narrowest distribution among the known subgenera of fiddler crabs. Its distribution restricted to Taiwan and the offshore Penghu Islands also implies the cladogenesis of this subgenus is related with the geological history of Taiwan Island during early Pliocene (reviewed by Shih et al. 2006, 2009b, 2011) and more studies are necessary to clarify it. Shih et al. (1999) have suggested this species to be included in the list of protected species of the Wildlife Conservation Law, R.O.C., because several habitats have been destroyed by industrial development, illegal fishing ponds, and mangrove overgrowth (Fig. 1; Shih et al. 1999). However, this species has not be considered as a protected species yet, although some populations of U. formosensis have been under different pressures, e. g. the Penghu population has become extinct in 2008 after the inappropriate replanting of introduced mangroves for about 15 years; and an unthoughtful ecological engineering on the habitat of Shengang, Changhua in 2006 nearly exterminated the local population (Shih 2008 a, b). The conservation policy and management for U. formosensis have to be reconsidered by government of Taiwan, because it is not only an endemic species to Taiwan, but also an endemic subgenus of fiddler crabs., Published as part of Shih, Hsi-Te, 2015, Uca (Xeruca), a new subgenus for the Taiwanese fiddler crab Uca formosensis Rathbun, 1921 (Crustacea: Decapoda: Ocypodidae), based on morphological and molecular evidence, pp. 151-169 in Zootaxa 3974 (2) on pages 162-165, DOI: 10.11646/zootaxa.3974.2.1, http://zenodo.org/record/242100, {"references":["Rathbun, M. J. (1921) New species of crabs from Formosa. Proceedings of the Biological Society of Washington, 34, 155 - 156.","Maki, M. & Tsuchiya, K. (1923) A Monograph of the Decapod Crustacea of Formosa. Report, Department of Agriculture, Government Research Institute, Taihoku 3. [in Japanese]","Barnwell, F. H. (1982) The prevalence of male right-handedness in the Indo-West Pacific fiddler crab Uca vocans (Linnaeus) and U. tetragonon (Herbst) (Decapoda: Ocypodidae). Journal of Crustacean Biology, 2, 70 - 83. http: // dx. doi. org / 10.2307 / 1548114","Shih, H. - T. (1988) A Preliminary Report of Crabs Collected in the Intertidal Zone at the Mouth of Tatu River (Crustacea: Decapoda). Bachelor's Thesis. Tunghai University, Taichung. [unkown pagination, in Chinese]","Fukui, Y., Wada, K. & Wang, C. - H. (1989) Ocypodidae, Mictyridae and Grapsidae (Crustacea: Brachyura) from some coasts of Taiwan. Journal of Taiwan Museum, 42, 225 - 238.","Huang, J. - F., Yu, H. - P. & Takeda, M. (1989) Fiddler crabs (Crustacea: Decapoda: Ocypodidae) of Taiwan. Bulletin of the Institute of Zoology, Academia Sinica, 28, 191 - 209.","Wang, C. - H. & Liu, H. - C. (1996 a) Common Seashore Crabs of Taiwan. Taiwan Museum, Taipei, 136 pp. [in Chinese]","Ho, P. - H. & Hung, M. - S. (1997) Seashore Crabs of Hsin Chu City. Hsinchu City Government, Hsinchu, 122 pp. [in Chinese]","Shih, H. - T., Mok, H. - K., Chang, H. - W. & Lee, S. - C. (1999) Morphology of Uca formosensis Rathbun, 1921 (Crustacea: Decapoda: Ocypodidae), an endemic fiddler crab from Taiwan, with notes on its ecology. Zoological Studies, 38, 164 - 177.","Hung, K. - H. (2000) Common Marine Biology in Penghu. Bureau of Culture, Penghu County, Penghu, 218 pp. [in Chinese]","Lee, J. - H. & Tung, S. - C. (2000) Crabs of Tainan County Coast. Tainan County Government and Kung San Institute of Technology, Tainan, 72 pp. [in Chinese]","Chen, Y. - H. (2001) Seashore Life (2): 700 Intertidal Species in Taiwan. Recreation Press, Taipei, 279 pp. [in Chinese]","Lee, J. - H. (2001) A Field Guide to Crabs in Taiwan. Bigtrees Co., Taipei, 174 pp. [in Chinese]","Rosenberg, M. S. (2001) The systematics and taxonomy of fiddler crabs: a phylogeny of the genus Uca. Journal of Crustacean Biology, 21, 839 - 869. http: // dx. doi. org / 10.1163 / 20021975 - 99990176","Shen, Y. - L. & Jeng, M. - S. (2005) Crabs of Penghu. Fisheries Research Institute, COA, Keelung, vi + 239 pp. [in Chinese]","Beinlich, B. & von Hagen, H. O. (2006) Materials for a more stable subdivision of the genus Uca Leach. Zoologische Mededelingen, 80, 9 - 32.","Wang, Y. - T. (2009) Intertidal Biota of Tainan. Tainan County Government, Tainan, 303 pp. [in Chinese]","Liu, H. - C. & Wang, C. - H. (2010) Taiwan Coastal Wetland Crabs. Wild Bird Society of Taipei, Taipei, 79 pp. [in Chinese]","Lee, J. - H., Chiu, Y. - W., Wu, Z. - Z., Wu, T. - T., Tseng, L. - K. & Huang, Y. - C. (2013) A Field Guide to the Shrimps, Crabs and Shells of Taijiang National Park. Taijiang National Park, Tainan, 239 pp. [in Chinese]","Sakai, T. (1939) Studies on the Crabs of Japan. IV. Brachygnatha, Brachyrhyncha. Yokendo, Tokyo, 377 pp. [pp. 365 - 741]","Horikawa, Y. (1940) A list of Formosan crabs. Scientific Taiwan 8, 21 - 31. [in Japanese]","Lin, C. - C. (1949) A catalogue of brachyurous Crustacea of Taiwan. Quarterly Journal of the Taiwan Museum, 2, 10 - 33.","Crane, J. (1975) Fiddler Crabs of the World (Ocypodidae: Genus Uca). Princeton University Press, Princeton, New Jersey, xxiii + 736 pp.","Sakai, T. (1976) Crabs of Japan and the Adjacent Seas. Kodansha Ltd., Tokyo, 773 pp.","Su, H. - J. & Lue, K. - Y. (1984). The crab fauna of Tan-shui mangrove swamp. Biological Bulletin of National Taiwan Normal University, 19, 61 - 70. [in Chinese]","Dai, A. - Y., Yang, S. - L., Song, Y. - Z. & Chen, G. - X. (1986) Crabs of the China Seas. China Ocean Press, Beijing, 642 pp. [in Chinese]","Dai, A. - Y. & Yang, S. - L. (1991) Crabs of the China Seas. China Ocean Press, Beijing, 682 pp.","Shih, H. - T. (1994) Fiddler Crabs. National Museum of Marine Biology / Aquarium, Pingtung, xiv + 190 pp. [in Chinese]","Ho, P. - H. (1996) Fiddler crabs (7): Uca formosensis. Fisheries Extension, Taipei, 116, 9 - 12. [in Chinese]","Shih, H. - T., Kamrani, E., Davie, P. J. F. & Liu, M. - Y. (2009 a) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U. lactea species-complex. Hydrobiologia, 635, 373 - 382. http: // dx. doi. org / 10.1007 / s 10750 - 009 - 9930 - 6","Shih, H. - T., Ng, P. K. L. & Liu, M. - Y. (2013 b) Systematics of the Indo-West Pacific broad-fronted fiddler crabs (Crustacea: Ocypodidae: genus Uca). Raffles Bulletin of Zoology, 61, 641 - 649.","Naderloo, R., Turkay, M. & Chen, H. - L. (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, 1 - 38.","Bott, R. (1973) Die verwandtschaftlichen Beziehungen der Uca Arten. Senckenbergiana Biologica, 54, 315 - 325.","Brosing, A. & Turkay, M. (2011) Gastric teeth of some thoracotreme crabs and their contribution to the brachyuran phylogeny. Journal of Morphology, 272, 1109 - 1115. http: // dx. doi. org / 10.1002 / jmor. 10967","Sakai, K., Turkay, M. & Yang, S. - L. (2006) Revision of the Helice / Chasmagnathus complex (Crustacea: Decapoda: Brachyura). Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 565, 1 - 76.","Brosing, A. (2010) Recent developments on the morphology of the brachyuran foregut ossicles and gastric teeth. Zootaxa, 2510, 1 - 44.","Shih, H. - T., Mok, H. - K. & Chang, H. - W. (2005) Chimney building by male Uca formosensis Rathbun, 1921 (Crustacea, Decapoda, Ocypodidae) after pairing: a new hypothesis for chimney function. Zoological Studies, 44, 242 - 251.","Shih, H. - T., Hung, H. - C., Schubart, C. D., Chen, C. A. & Chang, H. - W. (2006) Intraspecific genetic diversity of the endemic freshwater crab Candidiopotamon rathbunae (Decapoda, Brachyura, Potamidae) reflects five million years of geological history of Taiwan. Journal of Biogeography, 33, 980 - 989. http: // dx. doi. org / 10.1111 / j. 1365 - 2699.2006.01472. x","Shih, H. - T., Yeo, D. C. J. & Ng, P. K. L. (2009 b) The collision of the Indian plate with Asia: molecular evidence for its impact on the phylogeny of freshwater crabs (Brachyura: Potamidae). Journal of Biogeography, 36, 703 - 719. http: // dx. doi. org / 10.1111 / j. 1365 - 2699.2008.02024. x","Shih, H. - T., Ng, P. K. L., Naruse, T., Shokita, S. & Liu, M. - Y. (2011) Pleistocene speciation of freshwater crabs (Crustacea: Potamidae: Geothelphusa) from northern Taiwan and southern Ryukyus, as revealed by phylogenetic relationships. Zoologischer Anzeiger, 250, 457 - 471. http: // dx. doi. org / 10.1016 / j. jcz. 2011.07.004","Shih, H. - T. (2008 a) A preliminary study on the diversity and community composition of the fiddler crab [s] in Pong-Hu [Penghu]. Notes and Newsletter of Wildlifers, 12 (4), 17 - 21. [in Chinese]","Shih, H. - T. (2008 b) A Study on the Fiddler Crab, Uca formosensis, and Its Habitat. Hsinchu City Government, Hsinchu, 104 pp. [in Chinese]"]}

Sodhiana iranica n. sp. (Figs. 1���3) Material examined. Holotype: male (30.2 �� 23.1 mm) (ZRC 2014.0240), Bastak, Iran, 27 �� 13 ���N 54 �� 40 ���E, coll. S. Sharifian, 12 March 2014. Paratypes: 1 male (29.3 �� 23.0 mm), 3 females (largest 29.0 �� 23.1 mm) (ZRC 2014.0241), same data as holotype; 5 males (largest 30.3 �� 26.1 mm), 4 females (largest 33.8 �� 29.4 mm) (ZCUH), Bastak, Iran, 27 �� 13 ���N 54 �� 40 ���E, coll. S. Sharifian, 1 May 2014. Comparative material [photographs examined]. Sodhiana blanfordi (Alcock, 1909): Syntype male (46.4 �� 35.0 mm) (NHM 1905.10.13.5), Pishin, Baluchistan, Pakistan; syntypes, 4 males (largest 43.2 �� 33.2 mm), 5 females (largest 36.9 �� 28.8 mm) (ZSI 4043 / 4), Pishin, Baluchistan, Pakistan, coll. W.T. Blanford, no date; 1 male (41.8 �� 31.8 mm), 1 female (37.3 �� 28.8 mm) (ZRC), Punjgoor, Beluchistan, Pakistan, coll. Q. Kazmi, 19 January 2003; 2 males (larger 37.1 �� 28.0 mm) (ZRC), Punjgoor, Beluchistan, Pakistan, coll. Q. Kazmi, 19 August 2003. Description. Carapace broader than long; dorsal surfaces smooth; epigastric cristae low, formed by low transverse striae, not cristate, slightly anterior to postorbital cristae; postorbital cristae distinct, sharp, low, defined by small rounded granules, just reaching anterolateral margin at epibranchial tooth. External orbital tooth broadly triangular, outer margin lined with small rounded granules, about 3 times length of inner margin, margin near epibranchial tooth more angular, subdentiform. Epibranchial tooth distinct, triangular, just posterior of postorbital cristae, separated from margin by distinct V-shaped cleft (Figs. 1 A, B, 2 A, B). Third maxilliped ischium subrectangular, smooth, with submedian depression but no obvious longitudinal median sulcus; merus rectangular; exopod slender, reaching to midlength of merus, flagellum elongate (Fig. 2 C). Cutting edge of propodus of larger chela with large proximal tooth formed by fusion of several smaller teeth; tip of finger subspatulate (Fig. 2 F, H, I). Male thoracic sternum smooth; male abdomen narrowly triangular, somite 6 trapezoidal with very slightly concave lateral margins, longer than broad; longer than telson (Figs. 1 C, 2 D). G 1 with terminal segment straight, subconical, tip directed upwards, slightly inwards (toward median part of thoracic sternum), about one-fifth length of subterminal segment. G 2 distal segment long, about half length of basal segment (Fig. 3). Female vulvae rounded, on anterior half of sternite 6, with large opercular cover (Fig. 2 E). Habitat. The species inhabits a freshwater spring located in a semi-mountainous area in Iran (Eellod region), covered by dense stands of common reeds and salt cedar trees in the periphery, with mats of algae on the bottom. Distribution. So far only known from Bastak, Iran (but see ���Remarks��� below). Etymology. The specific epithet iranica is named after the type locality. Remarks. Sodhiana blanfordi (Alcock, 1909) and S. iranica n. sp. are superficially similar in the form of the carapace and ambulatory legs. The most significant difference is in the form of the G 1. The terminal segment of the G 1 of S. iranica n. sp. is gently bent inwards, with the outer margin more curved and the tip directed towards the median part of the sternoabdominal cavity (Fig. 3 A���C). In S. blanfordi, the terminal segment is evenly conical and straight (Fig. 4 F, G). These differences are consistent for the specimens of different sizes of both species studied. Sodhiana iranica n. sp. is easily separated from S. rokitanskyi by having the epigastric cristae lower and less pronounced, and coming close or almost in contact with the postorbital cristae (Figs. 1 A, 2 A, B), whereas it is more prominent in S. rokitanskyi (Pretzmann 1971: pl. 5 fig. 11; Kamrani et al. 2009: fig. 1 A; Yeo & Ng 2012: fig. 2 B). The G 1 of S. iranica n. sp. is relatively more slender with the terminal segment proportionately shorter and tip gently bent inwards (Fig. 3 A���C), in contrast to S. rokitanskyi, where the G 1 is proportionately stouter and the tip of the G 1 terminal segment is gently bent outwards (Pretzmann 1971: pl. 6 fig. 16; Kamrani et al. 2009: fig. 4 A, B). Sodhiana iranica n. sp. can be separated from the poorly known S. afghaniensis (Pretzmann, 1963) by its less convex dorsal carapace surface and proportionately longer G 1 terminal segment that has the tip directed inwards (versus straighter G 1 with a relatively shorter terminal segment in S. afghaniensis) (cf. Pretzmann 1963: pl. 2 figs. 5, 6; Yeo & Ng 2012: fig. 2 A). The carapace of the holotype and only specimen of S. afghaniensis is partially crushed but fresh material from near the type locality shows that adults have a distinctly convex dorsal carapace surface (P.K.L. Ng, unpublished). Kamrani et al. (2009) discussed the differences between S. rokitanskyi and S. blanfordi, and noted that the form of the posterior margin of the epistome and structure of the proximal tooth on the propodal finger of the larger chela were different. These characters may be affected by size. Although the specimens of S. blanfordi examined (through the courtesy of P.K.L. Ng) had the proximal tooth on the pollex of the larger chela composed of several smaller teeth, Alcock (1910: pl. 4 fig. 16 A) figured a large male that had a large complete tooth. This suggests that the large tooth is merely the consequence of complete fusion of several smaller teeth, certainly the case in the present material of S. iranica (Fig. 2 F, H, I). The same appears to be true for the form of the median triangular lobe on the posterior margin of the epistome. Smaller specimens have the lateral margins gently concave (cf. Kamrani et al. 2009: fig. 2 B) but larger ones have almost straight lateral margins (Fig. 4 C, D). It is noteworthy that the locality of the new species in Bastak, Iran, is over 1000 km to the west of the type locality of S. blanfordi (Pishin, Baluchistan) in western Pakistan, but separated by various high mountain ranges. The type locality of S. rokitanskyi occurs in near a hot spring in Geno, southern Iran (Kamrani et al. 2009), this area is some 200 km to the east of Bastak, Iran, the type locality of S. iranica n. sp., Published as part of Sharifian, Sana, Kamrani, Ehsan & Sharifian, Salim, 2014, A new species of freshwater crab of the genus Sodhiana Yeo & Ng, 2012 (Decapoda: Brachyura: Gecarcinucidae) from southern Iran, pp. 485-490 in Zootaxa 3878 (5) on pages 486-489, DOI: 10.11646/zootaxa.3878.5.7, http://zenodo.org/record/227753, {"references":["Alcock, A. (1909) Diagnoses of new species and varieties of freshwater crabs. Records of the Indian Museum, 4 (3), 375 - 381.","Pretzmann, G. (1971) Ergebnisse einiger Sammelreisen nach Vorderasien I. Teil: Zwei neue Unterarten von Susswasserkrabben. Annalen des Naturhistorischen Museums in Wien, 75, 473 - 475. [plates 1 - 6.]","Kamrani, E., Ng, P. K. L., Mirzadeh, M. & Nakhodai, S. (2009) The marsh crab, Sodhiana rokitanskyi (Pretzmann, 1971) (Decapoda, Brachyura, Gecarcinucidae) from southern Iran. Zootaxa, 2305, 24 - 32.","Yeo, D. C. J. & Ng, P. K. L. (2012) Sodhiana, A new genus of freshwater crab from south Asia (Decapoda: Brachyura: Gecarcinucidae). Raffles Bulletin of Zoology, 25, 279 - 284.","Pretzmann, G. (1963) Weiterer Bericht uber die mediterranen und vorderasiatischen Potamoniden. Annalen des Naturhistorischen Museums in Wien, 66, 373 - 380. [patels 1 - 3.]","Alcock, A. (1910) Brachyura I. Fasc. II. The Indian Freshwater Crabs- Potamonidae. Catalogue of the Indian Decapod Crustacea in the collection of the Indian Museum. Calcutta, 135 pp. [plates 1 - 14.]"]}

Uca (Austruca) albimana (Kossmann, 1877) (Figs. 1 a���l, 4 a, 6 a���b) Gelasimus annulipes var. albimana Kossmann, 1877: 53 ���54. Uca annulipes ��� Nobili 1906 a: 150 ���151; 1906 b: 311 (in key), 312. ��� Laurie 1915 (in list): 416. ��� Balss 1924: 15. ��� Hornby 1997: 15. Uca (Celuca) lactea annulipes ��� Crane 1975: 299, 301, 611. [part.: Red Sea material]. Uca (Celuca) lactea albimana ��� Lewinsohn 1977: 61 ���63. Uca lactea ��� Hogarth, 1989: 114 ���115. ��� Ismail & Ahmed 1993: 158 (in list). Uca lactea annulipes ��� Hywel-Davies 1994: 37, 48. Uca annulipes albimana ��� Apel & T��rkay 1999: 133. ��� Apel 2001: 114. Uca albimana ��� Shih et al. 2009: 377. Lectotype. 1 male [(SMF 9709) CL 6.8, CB 12.3, Ch.L 19.8 mm] Red Sea, 1874 - 75, leg. R. Kossmann. Paralectotypes. 1 male, 1 female (SMF 17145), data as lectotype. Other material. Red Sea: 2 males (ZMH K 2905), vend. Mus. Goddefroy; 1 male, 6 females (SMF 5698), Egypt, Kuseir, 0 5.1965, leg. E. Linsenmair; 1 male (SMF 26011) Egypt, Manteqat al Bahr al Ahmar, Safaga S. Hurghada, mangroves, 20.09. 1994, leg. C. Rhode & N. Dressler; 1 female (SMF 26012), Egypt, Manteqat al Bahr al Ahmar, 25 �� 52 'N 34 �� 25 'E, mangroves, 22.09. 1994, leg. C. Rhode & N. Dressler. Gulf of Aden: 1 male, 1 female (SMF 5646), Ethiopia, Massaua, 11.1826 ���02.1832, 1 eg. E. R��ppell; 1 male (MNHN B 11874), Yemen, Perim Island, 1897, leg. M. Jousseaume; 5 males (NHM 1974: 92) Jebel Hadd, near Aden, 10.1966, K. England; 1 male (MNHN B 11876), Djibouti, Obock, 1897, 1eg. M. Jousseaume; 216 males, 18 females (ZMH K 38114), Djibouti, winter 1909 /10, 1eg. E. Wache; 2 males (SMF 24497) Djibouti, Godoriyah, landward ridge of mangroves, 24.06. 1996, leg. U. Zajonz & F. Krupp. Arabian Sea: 3 males (NHM 1988: 55), Oman, Dhofar, 20 km N. of Ras Sawqirah, Khor Fakr, 18 �� 19 'N 56 �� 36 'E, 10.11.1985, 1eg. J.B. Clarke; 1 male (NHCY 83), Yemen, Sokotra, N. coast, Khawr Qariyah, 12 �� 38.054 'N 54 �� 12.568 'E, muddy-sand, 10.02. 1199, N. Simoes; 3 males, 1 female (NHCY 84), Yemen, Sokotra, W. coast, 12 �� 41.099 'N, 53 �� 28.476 'E, muddy-sand, bank of Wadi, 10.03. 1999, M. Apel; 2 males (NHCY 85), Yemen, Sokotra, bank of Khawr, 12 �� 35.976 'N 53 �� 46.44 'E, muddy, 0 4.02. 1999, N. Simoes; 3 males, 1 female (SMF 36907) Yemen, Sokotra, N. caost, Qadub-Mori, near Mori Lagune, 12 �� 38.446 'N, 53 �� 56.137 'E, muddysand, 05.02.1999; N. Simoes; 1 male (SMF 36908) Yemen, Sokotra, Khawr Qarayah, 12 �� 38.425 'N, 54 �� 13.338 'E, 0 7.04. 1999, M. Apel; 7 males, 2 females (1 ovig.), 1 juv. (SMF 36909), Yemen, Sokotra, Qualansiyah Lagune, 12 �� 41.904 'N, 53 �� 30.041 'E, muddy-sand, 12.04. 1999, M. Apel. Gulf of Oman: 13 males, 1 female (SMF 26013), UAE, Fujairah, Khor Kalba, 25 �� 01'N 56 �� 21 'E, mangroves, intertidal flat, sandy-mud substrate, 0 1.07. 1995, M. Apel; 1 male (SMF 26014), UAE, Fujairah, Khor Kalba, 25 �� 01'N 56 �� 21 'E, mangroves, intertidal flat, sandy-muddy substrate, 0 1.07. 1995, M. Apel. Persian Gulf: 3 males (SMF 26018), UAE, Ras al Khaimah, 25 �� 50 'N 55 �� 00'E, sandy flat, supralittoral zone among Salicornia sp., 11.07. 1995, M. Apel; 9 males (SMF 26015), UAE, Umm al Qaiwain, khor, 25 �� 35 'N 55 �� 34 'E, 0 9.07. 1995, M. Apel; 1 male (SMF 26032), UAE, Ajman, 25 �� 25 'N 55 �� 29 'E, intertidal flat, loose muddy substrate, among halophyte, 10.07. 1995, M. Apel; 1 male, 1 female (SMF 26017), UAE, Ras al Khaimah, mangroves of N. Rams, 25 �� 50 'N 55 �� 00'E, sandy-mud substrate, supralittoral zone, 10.07. 1995, M. Apel. Unknown locality: 3 males (MNHN), 0 2.1933, leg. J. L. Dantan. a, infraorbital region of male; b, infraorbital region of female; c, major chela, outer surface; d, major chela, inner surface; e, merus of major chela, outer surface; f, merus of major chela, inner surface; g, minor chela, outer surface; i, minor chela, outer surface; k, tip of G 1, mesial surface; j, tip of G 1, lateral surface. l, genital opening of female. Diagnosis. Front broad. Anterolateral borders of carapace moderately convergent. Major male chela palm without a supramarginal groove adjacent to lower border. G 1 with palp relatively short, not reaching to base of horny endpiece; this last one flat, with flanges not inclined, longer than broad, suture displaced ventrally; terminal opening marked by a more or less distinct notch. Redescription. Front broad, about 1 / 6.5 times as broad as carapace. Ventrolateral margin of carapace moderately convergent, distinctly crested, joining to posterolateral margin at base of exorbital triangle; posterolateral margin distinctly crested, extending curvedly backwards, ending at level of middle cardiac region; posterolateral striae short. Exorbital angle acutely triangular, directed forwards and inwardly; upper orbital margin sinuous, with two crests bordering narrow eyebrow; eyebrow getting narrower outwardly, inner part less than 1 / 2 breadth of eyestalk in adjacent region; lower orbital margin regularly granulate, granules becoming larger outwardly (Fig. 1 a), that of the paralectotype female with additional small granules, on middle part, at inner side of large granules (Fig. 1 b). Pterygostomian region (Fig. 1 a) with regular feather-shaped bristles, becoming glabrous towards lower orbital region, two short longitudinal lines of bristles on anterior-inner portion. Third maxilliped with merus about 1 / 3 length of ischium; longitudinal broad groove on outer surface of ischium and merus close to inner margin, bordered with short setae; long setae on inner margin of ischium and merus. Major chela merus with transverse ridges on upper margin, finely granulate, becoming irregular distally; lower margin denticulate (Fig. 1 e), denticles small, becoming larger distally; inner surface (Fig. 1 f) with patch of relatively large granules distally on upper portion. Carpus about 1.5 times as long as broad, inner upper margin denticulate, small patch of feather-shaped setae on lower proximal and upper distal portions of inner surface. Outer surface of palm (Fig. 1 c) smooth, without supramarginal groove; lower margin weakly granulate, with low granules, upper margin granulate, with relatively small granules; inner surface (Fig. 1 d) with high oblique tuberculate ridge, tubercles large, becoming smaller distally; proximal ridge at dactylus base with large granules, distal ridge parallel to proximal one, with granules, low, relatively small. Dactylus about 1.3 to 1.4 times as long as palm, higher than fixed finger; proximal, median teeth on cutting edge of both fingers, median teeth relatively large, that of fixed finger slightly larger; fixed finger with subdistal tooth. Small chela (Figs. 1 g, i) with smooth ridge on outer surface, extending from lower distal part of palm to two third of fixed finger; cutting edge without tooth, gap between fingers as wide as adjacent dactylus, fixed finger slightly wider than dactylus. Merus of walking legs with finely granulated transverse ridges on anterior margin, posterior margin minutely serrated; propodus 1.2 times as long as carpus; dactylus conical, as long as propodus; last leg with propodus as long as carpus; anterior margin of carpus and propodus slightly granulate. Male abdomen tapering from segment 2 to 5, segment 6 with lateral margin nearly straight, length of segments 3-5 subequal. G 1 (Figs. 1 j, k) with stem slightly bent in posterolateral direction, palp not reaching to base of horny endpiece; latter slightly longer than wide, distal margin convex, dorsal lobe large, reaching slightly beyond ventral lobe; terminal opening of sperm channel ventrally subdistal, in middle line of endpiece lobes; scarce feather-shaped setae along lateral margin, distally becoming longer and relatively dense, scarce short setae on palp. Genital opening of female: direction of opercle edge parallel to median line of sternum. Median tooth plate of the gastric mill (Fig. 4 a) with 8 teeth, small gaps between them, two first ones massive, different in shape, 3���8 decreasing in size posteriorly, last one distally attached to basal plate; lateral tooth plate with 20���22 comb-shaped teeth. Distribution. Red Sea, Gulf of Aden, Socotra, Arabian Sea, Oman (Dhofar), south Gulf of Oman, southeastern Persian Gulf. Remarks. Crane (1975), like former authors, treated the present species as a synonym of U. annulipes. Lewinsohn (1977) was the first to point out the differences between both and thus separated the Red Sea populations under the subspecific name U. lactea albimana. We agree with the differences cited by Lewinsohn (1977), namely the lack of a supramarginal groove on the major chela and lack of a patch of setae on the lower distal portion of merus of the same chela in all specimens from the Red Sea and specimens from outside the Red Sea. Furthermore, a distinct difference is the median tooth of the gastric mill. U. albimana has 8 teeth on the median plate, whereas U. iranica has 6, which confirms the clear difference between these two species. We therefore consider both of them as having species rank. Regarding the median tooth of the gastric mill, U. albimana is more closely related to U. annulipes, as this last species has also 8 teeth on the median tooth (Figs. 4 a, b). The most prominent difference is the shape of the second tooth, which is nearly elliptical in U. annulipes. The material from Oman, Gulf of Oman, Persian Gulf and M. Jousseaume's material from Djibouti (Obock) belong to the typical U. albimana. This species is remarkably small-sized (the largest male [SMF 5698] from the Red Sea measures CL 8.5, CB 15.5, Ch.L 25.00 mm) in comparison to its more closely related and partly sympatric congeners U. annulipes and U. iranica. Lewinsohn (1977) also mentioned the small size of U. albimana in comparison to U. annulipes. This species are sympatric, with U. iranica in the southeastern Persian Gulf and western Gulf of Oman, but easily distinguished from U. iranica, apart from its fully different median tooth of the gastric mill, by having moderately convergent lateral margins, which are very slightly convergent, or nearly straight in U. iranica. Genetic evidence presented by Shih et al. (2009) supports our morphological findings as to the differentiation of these two species from each other and from U. annulipes., Published as part of Naderloo, Reza, T��rkay, 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, pp. 1-38 in Zootaxa 2500 on pages 4-6, DOI: 10.5281/zenodo.195791, {"references":["Kossmann R. (1877) Malacostraca (1. Teil: Brachyura). Zoologische Ergebnisse einer im Auftrage der Koniglichen Academie der Wissenschaften u Berlin ausgefuhrten Reise in die Kustengebiete des Roten Meeres, 3 (1), 66 pp, pls 1 - 3.","Nobili G. (1906 a) Crustaces decapodes et stomatopodes. In: Mission J. Bonnier et Ch. Perez (Golfe Persique, 1901). Bulletin Scientifique de la France et de la Belgique, 40, 13 - 159, pls. 2 - 7.","Laurie R. D. (1915) Reports on the marine biology of the Sudanese Red Sea, XXI. On the Brachyura. Journal of the Linnean Society of London, Zoology, 31, 407 - 475, figs. 1 - 5, pls. 42 - 45.","Balss H. (1924) Decapoden des Rotes Meeres iii. Die Parthenopoiden. Cyclo- und Catometopen. Expedition S. M. Schiff ' Pola' in das Rote Meer, Nordliche und Sudliche Halfte, 1895 / 96 - 1897 / 98. Zoologische Ergebnise xxxiv. Denckschriften Akademie der Wissenschaften (Wien), 99, 1 - 18.","Hornby R. (1997) A survey of the habitats, invertebrate fauna and environmental sensitivity of the mainland coast of the UAE, with information on status and distribution of Crustaceans. TRIBULUS, Bulletin of the Emirates Natural History Group, 7 (2), 11 - 17.","Crane J. (1975) Fiddler Crabs of the World. Ocypodidae: Genus Uca. Princeton University Press, Princeton, New Jersey, i - xxiii + 736 pp.","Lewinsohn C. (1977) Die Ocypodidae des Roten Meeres (Crustacea, Decapoda, Brachyura). Zoologische Verhandelingen, 152, 45 - 84.","Hogarth P. J. (1989) The Marine Crustacea of Dhofar, Southern Oman. Journal of Oman Studies, 10, 99 - 124.","Ismail N. S. & Ahmed M. A. E. (1993) Macrobenthic invertebrates of mangrove, Avicennia marina (Forskal), and of intertidal flats of Khor Kalba, U. A. E., Gulf of Oman. In: Lieth, H. & Al Masoom, A. (Eds.), Towards The Rational Use Of High Salinity Tolerant Plants, Vol. 1. Tasks for Vegetation Sciences, 27 (1), 155 - 161.","Hywel-Davies A. (1994) A Quantitative Analysis of the Horizontal and Vertical Zonation of Brachyura and Mollusca Asociated with the Qurm Mangal, Muscat, Sultanate of Oman. MasterTthesis, School of Ocean Sciences, University of Wales, Bangor, 61 pp.","Apel M. & Turkay M. (1999) Taxonomic composition, distribution and zoogeographic relationships of the grapsid and ocypodid crab fauna of intertidal soft bottoms in the Arabian Gulf. Estuarine Coastal and Shelf Science, 49 (suppl. A), 131 - 142.","Apel M. (2001) Taxonomie und Zoogeographie der Brachyura, Paguridea und Porcellanidae (Crustacea: Decapoda) des Persisch-Arabischen Golfes. Unpublished Ph. D. Thesis, Johann Wolfgang Goethe-Universitat, Frankfurt am Main, 268 pp.","Shih H. - T, Kamrani E., Davie P. J. F & Liu M. Y. (2009) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U. lactea species-complex. Hydrobiologia, 635, 373 - 382."]}

Uca (Austruca) lactea (De Haan, 1835) (Figs. 10 a���f, 11 a���c, 14 a, 15 a, 18 a, b) Ocypode (Gelasimus) lactea De Haan 1835: 54, P 1. 15, fig. 5. Gelasimus lacteus ��� H. Milne Edwards 1852: 150, pl. 4, fig. 16. ��� Stimpson 1858: 100. ��� Miers 1879: 20, 36. ��� Kingsley 1880: 149, pl. 10, fig. 28. ��� Cano 1889: 234. ��� De Man 1891: 22. ��� Ortmann 1894 a: 752, 759. ��� Alcock 1900: 352 (in key), 355. ��� Lin 1949: 26. Uca lactea ��� Ortmann 1897: 351 ���355. ��� Balss 1922: 143. ��� Schenkel 1902: 580. ��� Gee 1925: 165. ��� Kellogg 1928: 356 (in list). ��� Gordon 1931: 528. ��� T. Sakai 1934: 320; 1936: 171; 1939: 618. ��� Kamita 1935: 241 (in list). ��� Miyake 1936: 511; 1939: 222, 242. ��� Ono 1965: 8. ��� Yamaguchi 1970: 5; 1973: 154; 1977: 426; 1978: 29; 2001 a: 285; 2001 b: 389. ��� Murai et al. 1987: 1334. ��� Jones & Morton 1994: 11 (in key), 28���30, fig. 6 A���I, Pl. 2 G, H, Pl. 3 E, F. ��� Iguchi et al. 1997: 70, fig. 2. ��� Yamaguchi & Ogata 2000: 133. ��� Ng et al. 2001: 37 (in list). ��� Tanaka et al. 2004: 8. ��� Kwok & Tang 2006: 4, figs. 10, 11. ��� Shih et al. 2009: 377. Austruca lactea ��� Bott 1973: 322, fig. 14. Uca (Celuca) lactea lactea ��� Crane 1975: 300, figs. 19 A, 54 J, 69 E, Pl. 40 A���B. ��� T. Sakai 1976: 608, P 1. 209, fig. 1. [part.]. ��� Miyake 1983: 162, pl. 55, fig. 2. Uca lactea lactea ��� Yamaguchi 1983: 25. ��� Kim & Kim 1995. ��� Shih et al. 1999: 170. Ucca (Celuca) lactea ��� Dai & Yang 1991: 466, fig. 236 (1), pl. 49 (2). Paralectotypes. 3 males, 1 female (ZMG 105), Japan, 1 eg. P. von Siebold. [exchanged with Leiden Museum in 1880]; 1 male (MNHN B 11967); 1 male (MNHN B 11964), [exchanged with Leiden Museum]. Other material. Korea: 2 males, 1 female (MNHN B 7912), Yellow Sea, Kanghwa-do, W of Seoul, 0 3.10. 1967, leg. H. S. Kim. Japan: 3 males (UZM); 5 males (ZMH K 3165), Shikoku, Sanuki, 13.03. 1897, ded. Leum; 1 male (ZSM), Okayama-ken, Okayama, freshwater, 1904 /05 1 eg. H. Sauter [Coll. Doflein]; 48 males, 2 females (ZMH K 2989), Kagoshima-ken, Satsuma Hanto, Tani Yama Wada, 29.12. 1898, vend. Faber & Voigt; 1 male (SMF 34991), Yoshinogawa River, 13.08. 1992, Hayashi & Hirata; 1 male (SMF 8183), Yoshinogawa River, 13.08. 1992, Hayashi & Hirata; 1 female (SMF 34968), Tokushima city, Sumiyoshi, Yoshinogawa River, 30.07. 1992, Hayashi & Hirata, ded. K. Sakai; 1 female (SMF 34969), Tokushima City, Sumiyoshi, Yoshinogawa River, 30.07. 1992, Hayashi & Hirata, ded. K. Sakai; 1 male (SMF 34999), Tokushima City, Sumiyoshi, Yoshinogawa River, 0 6.09. 1992, ded. K. Sakai; 5 males, 1 female (SMF 35003), Tokushima, Jotocho, mouth of the Yoshinogawa River, 0 9.08. 1983, K. Sakai; 1 male (SMF 35005), Tokushima, Ohjincho, Yoshinogawa River, 26.04. 1983, K. Sakai; 2 males (SMF 35006), Tokushima, Ohjincho, Yoshinogawa River, 0 5.07. 1981, K. Sakai; 2 males (SMF 35007), Kochi, Tosa Bay, Utsuga, 24.07. 198026.07. 1980, K. Sakai; 5 males, 2 females (SMF 35008), Hiroshima, Urazaki-cho, Onomichi, 02.09.1981 - 05.09.1981, Y. Kuwada, ded. K Sakai; 1 male, 1 female (SMF 35009), Kochi, Shimantogawa River, 0 1.08. 1996, K. Sakai; 5 males, 2 females (SMF 35010), Amakusa Island, Kyushu, 04.1974���05.1981, Y. Ono, ded. K Sakai; 1 male, 1 female (SMF 35011), Kochi, Irino, Okata-cho 0 1.08. 1996, K. Sakai. Taiwan: 1 male (SMF 8813) W. coast, Mai Liao, 23 �� 47 'N, 120 �� 11 'E, sandy-mud flat, 0 2.08. 1977, leg. J. D��rjes; 1 male, 2 females (juv.) (SMF 8814), 0 7.09. 1977, leg. J. D��rjes; 1 male (UZM), Tanshui River NW. of Taipei [ex Mus.Milano, 20.02.1922]; 2 Chelas (ZSM), Amping, 1 eg. Haberer; 1 male (ZSM), Takao, 0 6.1903, 1 eg. Haberer. China: 26 males, 5 females (ZMH K 6755); 2 males (MNHN B 11963); 2 males (ZSM); 3 males (MNHN B 11966), leg. M. Callery; 10 males, 3 females (ZMH Ex K 3100), 04.09.1919; 13 males, 3 females (ZMH ex K 3156), 28.10. 1910, leg. G. Siemssen; 1 male (IOASQ), Shajiang, 0 3.03.1957, 1eg. Fan & Xu; 4 males, 1 female (ZMH Ex K 3154), Fuzhou, 13.06. 1903, leg. G. Siemssen; 2 males (IOASQ), Quanzhou, Shengpu, 21.06.1975, 1eg. H.L. Chen; 2 males, 2 females (IOASQ), Tongan, Xiaodeng, N. of Xiamen, 0 2.04.1982, 1eg. H.L. Chen; 5 males, 1 female (MNHN B 11965), Xiamen, 0 7.1925, 1 eg. C.F. Wang; 2 males, 2 females (SMF 17158), Xiamen, Zhongzhai, 26.05. 1982, leg. H.L. Chen; 1 male, 1 female (SMF 17159), Dongshan, Beinanao, 26.05.1975, 1eg. H.L. Chen; 1 male (IOASQ); 2 males, 1 female (SMF 17160), Hainan, Haikou, Cangtonqian, 0 7.06. 1958, leg. J.Y. Liu; 1 male, 1 female (IOASQ); 2 males (SMF 17162), Haikou, Beigang, 0 6.06. 1958, leg. J.Y. Liu; 1 male (SMF 17161), Beili, 12.07.1957, 1eg. J.Y. Liu. Diagnosis. Front broad. Anterolateral borders moderately convergent. palm of major male chela without supramarginal groove adjacent to lower border. G 1 (Fig. 10 f) with palp long, reaching up to distal half of horny endpiece; this last one has inclined flanges, but no strong torsion (Fig. 11 c); terminal opening at base of deep emargination of distal margin (Fig. 11 b). Genital opening of female (Fig. 11 a) and thus also direction of the opercle's edge slightly oblique in respect to median line of sternum. Distribution. Korea, Japan, Taiwan, China, northern Vietnam. The northern limit in China is in Zhejiang Province, Ise Bay in Japan. Southwards the species reaches Hainan Island, China and northern Vietnam. Remarks. The species is easily recognisable by the peculiar shape of the G 1, especially the large size of the palp. U. lactea may easily be distinguished from U. perplexa, with which it may be sympatric at least in the Ryukyu Islands, by the lack of a supramarginal groove on the major chela palm as well as by the weakness of the subdistal tooth on the fixed finger of the same limb (Figs. 10 b���e), which is more prominent in U. perplexa and makes it easily distinguishable from its congener. The southern distribution of this species is not well known. This applies especially to the Ryukyus, from which it was reported by T. Sakai (1976). Although Sakai mentioned the presence of U. lactea in the Ryukyus, but he did not list U. perplexa from Japan. Sakai probably ignored Crane's record of U. perplexa in the Ryukyus. Crane (1975), however, only recorded U. perplexa from this area and our material also belongs to this species. On the contrary, we have not seen any specimen of U. perplexa from China. The same applies to our few specimens from Taiwan, which all belongs to U. lactea, although the species has been recorded from Taiwan several times (see Ng et al. 2001). It has to be checked if U. perplexa and U. lactea exclude each other or if they are sympatric in part of their range. The factors limiting the distribution of the two species also remain unknown. The median tooth of the gastric mill of U. lactea (Fig. 14 a) is unique among all member of the U. lactea- group, but slightly similar to U. perplexa rather than others. But regarding the form of the lateral tooth plate of the gastric mill (Fig. 15 a) and the number of teeth on it, numbering 22 and being comb-shaped, ally the species with U. annulipes and U. albimana., Published as part of Naderloo, Reza, T��rkay, 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, pp. 1-38 in Zootaxa 2500 on pages 19-21, DOI: 10.5281/zenodo.195791, {"references":["Milne Edwards H. (1852) Observation sur les affinites zoologiques et la classification naturelle des crustaces. Annales des Sciences Naturelles (Zoologie), (3) 18, 109 - 166.","Stimpson W. (1858) Prodromus descriptionis animalium evertebratorum, quae in Expeditione ad Oceanum Pacificum Septentrionalem, a Republica Federata missa, Cadwaladaro Ringgold et Johanne Rodgers Ducibus, observavit et descripsit. Pars V. Crustacea Ocypodoidea. Proceedings of the Academy of Natural Sciences of Philadelphia, 10, 93 - 110.","Miers E. J. (1879) On a collection of Crustacea made by Capt. H. C. St. John, R. N., in the Korean and Japanese seas. Part i. Podophthalmia. Proceedings of the Zoological Society of London, 1879, 17 - 61, 3 pls.","Kingsley J. S. (1880) Carcinological notes. Proceedings of the Academy of Natural Sciences of Philadelphia. For 1879: 34 - 37; ii. Revision of the Gelasimi, 135 - 152.","Cano G. (1889) Crostacei brachiuri ed anornuri raccolti nel viaggio della \" Vettor Pisani \" intorno al globo. Bolettino della Societa dei Naturalisti in Napoli, 3, 79 - 106, 169 - 269.","Man J. G. De (1891) Carcinological studies in the Leyden Museum, No 5. Notes from the Leyden Museum, 13, pp. 1 - 61.","Ortmann A. E. (1894 a) Die Decapoden-Krebse des Strassburger Museums. viii. Theil. Abteilung: Brachyura (Brachyura genuina Boas) iii. Unterabteilung: Cancroidea, 2. Section: Cancrinea, 2. Gruppe: Catometopa. Zoologische Jahrbucher, Abtheilung fur Systematik, Geographie und Biologie der Thiere, 7, 683 - 772, pl. 23.","Alcock A. (1900) Material for a carcinological fauna of India. No. 6: The Brachyura Catometopa, or Grapsoidea. Journal of the Asiatic Society of Bengal, 69 (2), 279 - 456.","Lin C. - C. (1949) A catalogue of brachyurous Crustacea of Taiwan. Quarterly Journal of the Taiwan Museum, 2, 10 - 33.","Ortmann A. E. (1897) Carcinologische Studien. Zoologische Jahrbucher. Abteilung fur Systematik, Geographie und Biologie der Thiere, 10 (3), 258 - 372.","Balss H. (1922) Ostasiatische Decapoden. iv. Die Brachyuren (Canceridae). Archive Fur Naturgeschichte, A (11), 94 - 166.","Schenkel E. (1902) Beitrag zur Kentnis der Dekapodenfauna von Celebes. Verhandlungen der Schweizerischen Naturforschenden Geselschaft (Basel, 13, 485 - 585. pls. 7 - 13.","Gee N. G. (1925) Tentative list of Chinese decapod Crustacea including those represented in the collection of the U. S. N. M. with localities at which collected. Lingnaam Agricultural Review, 3 (2), 156 - 66.","Kellogg C. R. (1928) Crustacea of Fukien Province. Lingnaan Science Journal, 5 (4), 351 - 356.","Gordon I. (1931) Brachyura from the coasts of China. Journal of the Linnean Society of London, Zoology, 37 (254), 525 - 558.","Sakai T. (1934) Brachyura from the coast of Kyushu, Japan. Science Reports of the Tokyo Bunrika Daigaku Section, 1 (25), 281 - 330.","Kamita T. (1935) On the Brachyura of the west Korean waters (Yellow Sea). Zoological Magazine (Tokyo), 47, 232 - 245.","Miyake S. (1936) A list of known species of Brachyura from Ishigakishima. Annotationes Zoologicae Japonenses, 15 (4), 506 - 513.","Ono Y. (1965) On the ecological distribution of ocypodoid crabs in the estuary. Memoirs of the Faculty of Science of Kyushu University, Series E, 4 (1), pp. 1 - 60.","Yamaguchi T. (1970) Ecology of a fiddler crab (Uca lactea). Calanus, 2, 5 30.","Murai M., Goshima S. & Henmi Y. (1987) Analysis of the mating system of the fiddler crab, Uca lactea. Animal Behaviour, 35 (5), 1334 1342.","Jones D. S. & Morton B. (1994) The fiddler crabs (Ocypodidae: Uca) of Hong Kong. Asian Marine Bailogy, 11, 9 - 40.","Iguchi R., Tajima M. & Wada K. (1997) Distribution of fiddler crabs, Uca arcuata and Uca lactea in the Yoshino River Estuary and vicinities (Tokushima Prefecture). Bulletin of the Tokushima Prefectural Museum, 7, 69 79.","Yamaguchi T. & Ogata R. (2000) Studies of the first and second maxillipeds of the fiddler crab, Uca lactea. Crustacean Research, 29, 133 142.","Tanaka H., Shibagaki K., Ikezawa H., Kanesawa A. & Wada K. (2004) Occurrence of two species of fiddler crab (Decapoda, Ocypodidae) on the tidal flat of the Aono River, Izu Peninsula, Shizuoka Prefecture, Japan. Japanese Journal of Benthology, 59, 8 12.","Kwok W. P. W. & Tang W. - S. (2006) Fiddler crabs in Hong Kong-an overview. Hong Kong Biodiversity, 12, 1 - 7.","Shih H. - T, Kamrani E., Davie P. J. F & Liu M. Y. (2009) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U. lactea species-complex. Hydrobiologia, 635, 373 - 382.","Bott R. (1973) Die Verwandtschaftliche Beziehungen de Uca - Arten (Decapoda: Ocypodidae). Senckenbergiana Biologica, 54, 315 - 325.","Crane J. (1975) Fiddler Crabs of the World. Ocypodidae: Genus Uca. Princeton University Press, Princeton, New Jersey, i - xxiii + 736 pp.","Sakai, T. (1976) Crabs of Japan and the Adjacent Seas. Kodansha, Tokyo, vol. 1 [English text], xxix + 773 pp., figs. 1 - 379, maps 1 - 3; vol. 2 [Japanese text], 461 pp., figs. 1, 2; vol. 3 [plates], 16 pp. + pls. 1 - 251.","Miyake S. (1983) Japanese crustacean decapods and stomatopods in color. 1. Macrura, Anomura and Stomatopoda. Osaka: Hoikusha. 261 pp.","Yamaguchi T. (1983) Representative shore animals in the Amakusa Islands (1). The fiddler crab, Uca lactea lactea (de Haan). Calanus, 8, 25 32.","Kim W. & Kim S. H. (1995) The brachyuran crabs (Crustacea: Decapoda) of Chindo Island, Korea. Korean Journal of Systematic Zoology, 11 (4), 497 508.","Shih H. - T, Mok H. - K, Chang H. - W & Lee S. - C. (1999) Morphology of Uca formosensis Rathbun, 1921 (Crustacea: Decapoda: Ocypodidae), an endemic fiddler crab from Taiwan, with notes on its ecology. Zoological Studies, 38 (2), 164 - 177.","Dai Ai-Yun & Yang S. (1991) Crabs of the China Seas. China Ocean Press Beijing; Springer Berlin, Heidelberg, New York, Tokyo, 682 pp., pls. 1 - 74 (English edition)."]}

Uca (Austruca) mjoebergi Rathbun, 1924 (Figs. 12 d���f, 13 a���j, 14 b, 15 b) Uca mjobergi Rathbun 1924: 9. Uca lactea mjoebergi ��� Crane 1975: 299, figs. 17 A���C, 19 O���P, 54 H, 69 B. Uca mjoebergi ��� George & Jones 1982: 10 (in key), 80���83, figs. 4 b, 7 a, 53 a���f, 58 d. ��� von Hagen & Jones 1986: 57 (in key), 66. ��� Morgan 1990: 62, 69 (in list). ��� Davie 2002: 362. ��� Shih et al. 2009: 377. ��� Barnes 2010: 250, fig. 1. Material examined. Indonesia: 1 male (NNM 32589), Indonesia, Irian Jaya (= Western New Guinea), mangroves near Kampong Madong, between Mokmer and Bosnek, SE Biak, 20.02. 1955, leg. L.B. Holthuis; 34 males, 14 females (UZM), 1 male (SMF 17163), Banda-Isles, Lontor, Coast near Lavvande, 0 6.06. 1922, "Dan. Expedition to the Kei Islands, 1922 "; 2 males, 1 female (UZM), same locality and expedition, 0 7.06. 1922. Australia: Northern Territory: 2 males (NHMB 574 C), Port Darwin, 1 eg. E. Handschin. ���Western Australia: 10 males, 13 females (ZMH), 2 males, 2 females (SMF 17155), Broome, mangroves, 24.11. 1975, leg. G. Hartmann & G. Hartmann-Schr��der; 10 males, 2 females (ZMH); 2 males (SMF 17156), Harbour, 1 km E. of Onslow, about 500 m. from mouth of Beadon Creek, 0 7.06. 1975, leg. G. Hartmannn & G. Hartmann- Schr��der; 12 males, 4 females (ZMH), Exmouth, external side of peninsula, Reef near Tantabiddy Creek, 11.10. 1975 or Carnavon, Pelican Point, inner beach with mangroves, 13.10. 1975, leg. G. Hartmann & G. Hartmann-Schr��der. Western Samoa: 1 male (SMF 5676), Upolu, vend. Mus. Goddefroy [this locality is probably wrong]. Diagnosis. Front broad. Anterolateral borders of carapace slightly convergent. Palm of major male chela (Figs. 13 c, d) without supramarginal groove adjacent to lower margin; oblique ridge at inner surface low. G 1 generally as in U. annulipes, but horny endpiece comparatively broader, directed remarkably in a dorsolateral direction (Fig. 13 e), mesial border of stem between palp and endpiece bulging outward. Distribution. Australia: Northern Territory and Western Australia, New Guinea (where it probably is sympatric with U. annulipes), central Indonesia (Sulawesi Tenggara, Pulau Kaledupa) sympatric with U. perplexa (Barnes 2010). Remarks. The most reliable characters to recognise this species is the shape of the G 1, especially the curb in the distal part of stem adjacent to the horny endpiece as well as its convexity, of course with a certain amount of variability (see Figs. 11 f���i). It is easily distinguished from U. perplexa by the lack of the supramarginal groove at the outer surface of the palm of the large male chela. From U. annulipes it can be also easily distinguished by this last character. All Australian specimens examined are typical both in the shape of the G 1 and the palm of the male chela. In the examined specimens also the oblique ridge on the inner surface of large male chela palm is typically low as described by Crane (1975) and George & Jones (1982). In U. annulipes this crest is typically high, but there is a certain amount of variability in this character, so that it cannot be used as good discrimination character. In the specimens from the Banda Archipelago different forms of G 1 were found, exhibiting some variations from typical U. annulipes to typical U. mjoebergi (see Fig. 11 f���i). The specimen from New Guinea identified by Crane (1975) as U. mjoebergi is quite typical in the major chela. Regarding the features of median and lateral teeth of the gastric mill, this species is readily separated from all other members of this group. The median tooth plate (Fig. 14 b) has four teeth, the two first ones being variable in size, the third and fourth ones nearly equal, one very small tooth attached dorsally to the basal plate. The lateral tooth plate (Fig. 15 b) is typical in having a shoe-shaped form, with 17 combshaped teeth. U. mjoebergi has a local distribution in northern and Western Australia (Davie 2002). There are also specimens from western New Guinea, where it is likely sympatric with U. annulipes. More recently, Barnes (2010) recorded this species from Sulawesi Tenggara in central Indonesia in sympatry with four different species of fiddler crabs, including U. perplexa from the present group. These records from outside Australia extend its distiribution north to South East Asia, but still within Wallacea., Published as part of Naderloo, Reza, T��rkay, 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, pp. 1-38 in Zootaxa 2500 on page 22, DOI: 10.5281/zenodo.195791, {"references":["Rathbun M. G. (1924) Results of Dr. E. Mjoberg's Swedish scientific expeditions to Australia 1910 - 1913. 37. Brachyura, Albuneidae and Porcellanidae. Arkiv for Zoologi, 16 (23), 1 - 33.","Crane J. (1975) Fiddler Crabs of the World. Ocypodidae: Genus Uca. Princeton University Press, Princeton, New Jersey, i - xxiii + 736 pp.","George R. W. & Jones D. S. (1982) A revision of the fiddler crabs of Australia (Ocypodidae: Uca). Record of the Western Australian Museum, Suppl. 14, 99 pp.","Jones D. A. (1986) A Field Guide to the Sea Shores of Kuwait and the Arabian Gulf. University of Kuwait Blandford Press, Poole, Kuwait (1986), 192 pp.","Morgan G. J. (1990) A collection of Thalassinidea, Anomura and Brachyura (Crustacea: Decapoda) from the Kimberley Region of northwestern Australia. Zoologische Verhandelingen, Leiden, 265, 90 pp, figs. 1 - 6.","Shih H. - T, Kamrani E., Davie P. J. F & Liu M. Y. (2009) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U. lactea species-complex. Hydrobiologia, 635, 373 - 382.","Barnes R. S. K. (2010) A remarkable case of fiddler crab, Uca spp., alpha diversity of Wallacea. Hydrobiologia, 637, 249 - 253."]}

Uca typhoni Crane, 1975 (Figs. 4 A, B, 5 A, C) Uca (Deltuca) demani typhoni Crane, 1975: 41, pl. 4 E���H, fig. 61 C [type locality Manila, Luzon]. Uca (Tubuca) typhoni ���Ng et al. 2008: 242 [list]. Material examined. Hainan: 1 �� (30.75 mm) (NCHUZOOL 13371), Sanya, Hainan, coll. H.-T. Shih & R.- H. Lee, 28 Jun. 2004; 1 �� (20.64 mm) (MBMCAS 166269), Qinglan Harbor, Wenchang, 5 Dec. 1990. Comparative material. Uca demani Ortmann, 1897: 2 3 (16.70, 20.54 mm), 3 ���� (17.92���26.53 mm) (NCHUZOOL 13372), Cebu, Philippines, coll. L. Liao et al., 2 Sep. 2003; 1 3 (19.09 mm) (NCHUZOOL 13373), Zamboanga, Philippines; 2 3 (27.45, 28.04 mm) (ZRC 2000.1731), Sulawesi, Indonesia, coll. C.D. Schubart, 19 Jan. 2000. Live coloration. The Chinese female specimen (NCHUZOOL 13371) has purple carapace and legs, except that two legs and the left minor cheliped are bright red (Fig. 4 A, B). This asymmetry is anomalous, and may be the result of these appendages having been lost and regenerated. Ecology. The Chinese specimen (NCHUZOOL 13371) was collected from the mangrove area at the southern tip of Hainan Island (Fig. 4 C). It occurred sympatrically with other species including Uca crassipes, U. annulipes, U. vocans, Mictyris brevidactylus Stimpson, 1858, and Macrophthalmus sp. (see Shih et al. 2009, 2010; Davie et al. 2010). Distribution. Northern Philippines and Hainan in China. It is a new record both for China and East Asia. Remarks. Crane (1975) reported three subspecies under U. demani, viz. U. demani demani Ortmann, 1897, U. demani typhoni Crane, 1975, and U. demani australiae Crane, 1975. Uca australiae was published based on a single specimen from Broome, Australia (Crane 1975) and its taxonomic validity is considered doubtful (George & Jones 1982). Crane (1975) identified the specimens from northern Philippines (Negros, Panay, Lubang, Samar, and Luzon) as U. typhoni (type locality Manila), and specimens from southern Philippines (Zamboanga and Gulf of Davao in Mindanao) and Indonesia (Java, Sumbawa, and Sulawesi) as U. demani (lectotype locality Sulawesi). For this study, we compared specimens from Hainan, Cebu (next to Negros), Zamboanga, and Sulawesi (see Comparative material). The specimens from southern Philippines and Indonesia agree well with the description of U. demani, especially with regard to the form of the male major cheliped (Crane 1975: 39; Yamaguchi 1994: fig. 4). The freshly preserved specimens show the reddish purple on the posterior carapace, and the degrees of gray blotching on the anterior part, and the gray legs (Fig. 4 E, F), conform well with Crane���s (1975: 42) description. Crane (1975: 41���42) mentioned the relative development of tubercles on the floor of the orbit is important to separate the three subspecies, e.g., tubercles of U. demani typhoni are 12���18 and U. demani demani 18���22. However, we found the number of tubercles on the orbital floor to be variable and unreliable. The tubercles on U. demani are 0���24 for males and 5���18 for females (cf. Fig. 5 B), and large specimens tend to have more tubercles, although the tubercles are weak or lacking on either side of the orbits for some specimens (Table 2). The females from Hainan have 16���28 tubercles on the orbits (Fig. 5 A), a similar number or more than those from Philippines and Indonesia (Table 2). The large variation of the tubercle number on the orbital floor makes this character not very unsuitable for separating these species. However, there are substantial differences in the gonopore morphology between the females from Hainan and Cebu (Fig. 5 C, D). Although Crane (1975: 40) noted that the subspecies of U. demani can be differentiated by the form of the gonopores, she also admitted that the ���[Female] Material is insufficient for a satisfactory comparison in either d. demani or d. typhoni ���. For the gonopore of the Hainan specimen, the depression is deep, with lateral-external margin surrounded by a low raised rim, and the angle of the lateral margin is subparallel to the longitudinal line (Fig. 5 C). For the gonopore of U. demani from Cebu, the depression is shallow with a median tubercle, the anterior and posterior rims are relatively more prominent and more raised, with the angle of the lateral margin directed slightly towards the postero-external part (Fig. 5 D). tubercle number on floor species sex CW left orbit right orbit locality museum cat. no. Uca typhoni �� 20.64 16 16 Hainan MBMCAS 166269 �� 30.75 28 26 Hainan NCHUZOOL 13371 Uca demani 3 16.70 1 2 Cebu NCHUZOOL 13372 3 19.09 13 16 Zamboanga NCHUZOOL 13373 3 20.54 7 8 Cebu NCHUZOOL 13372 3 27.45 13 24 Sulawesi ZRC 2000.1731 3 28.04 0 13 Sulawesi ZRC 2000.1731 �� 17.92 5 6 Cebu NCHUZOOL 13372 �� 22.54 16 17 Cebu NCHUZOOL 13372 �� 26.53 17 18 Cebu NCHUZOOL 13372 In addition, the genetic differences also support their separation as species. The Philippine and Indonesian specimens form a distinct clade, while the Hainan specimen belongs to another clade, with the basepair differences 1.9 % of 16 S rRNA and 4.1 % of COI (Shih et al. unpublished data). In conclusion, based on the morphology, distribution, and genetic data, we are confident that the female specimens from Hainan are U. typhoni. The occurrence of U. typhoni in southern Hainan extends its range westwards. It is also a new record for China and East Asia., Published as part of Shih, Hsi-Te, Ng, Peter K. L., Fang, Shou-Hua, Chan, Benny K. K. & Wong, Kingsley J. H., 2010, Diversity and distribution of fiddler crabs (Brachyura: Ocypodidae: Uca) from China, with new records from Hainan Island in the South China Sea, pp. 1-19 in Zootaxa 2640 on pages 13-15, DOI: 10.5281/zenodo.198567, {"references":["Crane, J. (1975) Fiddler Crabs of the World (Ocypodidae: Genus Uca). Princeton University Press, Princeton, New Jersey.","Ortmann, A. E. (1897) Carcinologische Studien. Zoologische Jahrbucher, Abtheilung fur Systematik, Geographie und Biologie der Thiere, 10, 258 - 372.","Stimpson, W. (1858) Crustacea Ocypodoidea. Prodromus descriptionis animalium evertebratorum, quae in Expeditione ad Oceanum Pacificum Septentrionalem a Republica Federata missa, Cadwaladaro Ringgold et Johanne Rodgers ducibus, observavit et descripsit, Pars 5. Proceedings of the Academia of Natural Science, Philadelphia, 10, 93 - 110.","Shih, H. - T., Kamrani, E., Davie, P. J. F. & Liu, M. - Y. (2009) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U. lactea species-complex. Hydrobiologia, 635, 373 - 382.","Davie, P. J. F., Shih, H. - T. & Chan, B. K. K. (2010) A new species of Mictyris (Decapoda, Brachyura, Mictyridae) from the Ryukyu Islands, Japan. Crustaceana Monographs, 11, 83 - 105.","George, R. W. & Jones, D. S. (1982) A revision of the fiddler crabs of Australia (Ocypodinae: Uca). Records of the Western Australian Museum, Supplement, 14, 5 - 99.","Yamaguchi, T. (1994) Fiddler crabs of the genus Uca in the collections of three natural history museums in Europe. 1. The specimens held by the Nationaal Natuurhistorisch Museum, Leiden and the Natural History Museum, London. Calanus, 11, 151 - 189."]}

Uca lactea (De Haan, 1835) Ocypode (Gelasimus) lactea De Haan, 1835: 54, pl. 15, fig. 5 [type locality Japan]. Gelasimus lacteus ���H. Milne Edwards 1852: 150 (not pl. 4 (16)) (Macao?); Stimpson 1858: 100; 1907: 108 [Macao]; Kingsley 1880: 149, pl. 10 (28) (part); Cano 1889: 92, 234 [Xiamen]; Shen 1940: 231 [Hong Kong]. Uca lactea ��� Balss 1922: 143 [list]; Gee 1926: 165 [Xiamen; Chin Bey; Hong Kong; Macao]; Kellogg 1928: 356 [list]; Gordon 1931: 528 [list]; Shen, 1936: 76 [Hainan]; Jones & Morton 1994: 28, pls. 2 G, H, 3 E, F, fig. 6 [Hong Kong]; Yamaguchi 1994: 165 [Xiamen]; Cai et al. 1998: 117 [Hong Kong]; Ying et al. 1999: 34 [Zhejiang]; Xiao & Li 2003: 61 [Naozhou, Zhanjiang, Guangdong]; Kwok & Tang 2006: 4, figs. 10, 11 [Hong Kong]; Yang et al. 2008: 807 [East China Sea; northern South China Sea]. Uca lacteus ��� Shen & Dai 1964: 111, unnumbered text-fig. [Fujian; Hainan]; Yang & Liu 1996: 189 [Zhanjiang, Guangdong]. Uca (Celuca) lactea lactea ��� Crane 1975: 300, pl. 40 A, B, figs. 19 A, 54 J-JJ, 69 E [Xiamen; Hong Kong]; Dai et al. 1986: 425, pl. 59 (2), fig. 236 (1) [Fujian; Hainan]. Uca (Celuca) lactea ��� Dai & Yang 1991: 466, pl. 59 (2), fig. 236 (1) [Fujian; Hainan]; Lu 1996: 165 [Xiamen, Fujian]; Lai & He 1998: 172 [Guangxi]; Liang et al. 2005: 22 [Leizhou, Guangdong]. Uca lactea lactea ��� Li 2004: 21 [Fujian]; Chen 1991: 411, fig. 388 [Zhejiang]. Uca (Paraleptuca) lactea ���Ng et al. 2008: 241 [list]. Uca (Austruca) lactea ��� Naderloo et al. 2010: 19, figs. 10 a���f, 11 a���c, 14 a, 15 a, 18 a, b [Fujian; Hainan]. Material examined. Fujian: 2 3 (19.5 mm, one damaged) (MBMCAS 166306), Xiunpu, Quanzhou, coll. H.-L. Chen, 21 Jun. 1975; 7 3 (6.89���15.97 mm), 1 �� (12.15 mm) (NCHUZOOL 13374), Jimei, Xiamen, coll. H.-T. Shih & J.-H. Lee, 1 Jul. 2004. Hong Kong: 5 3 (11.96���16.37 mm), 3 ���� (11.16���13.14 mm) (NCHUZOOL 13250), Tai Tam, coll. B. Chan, Jul. 2004; 1 3 (12.14 mm), 1 �� (12.32 mm) (NCHUZOOL 13375), Tai Tam, coll. B. Chan, 27 Oct. 2008. Hainan: 5 3 (13.26���14.14 mm), 3 ���� (10.82���11.87 mm) (NCHUZOOL 13376), Dongzhai, coll. H.-T. Shih & J.-H. Lee, 23 Jun. 2004. Guangxi: 7 3 (11.25���20.02 mm), 1 �� (10.9 mm) (NCHUZOOL 13377), Qinzhou, coll. H.-T. Shih & Y.-H. Wang, 10 May 2009. Distribution. Japan, Korea, China, Taiwan, and Vietnam (cf. Table 1). Remarks. Uca lactea is one of the most common species in East Asia and Vietnam. It ranges from Korea (northernmost distribution), Japan, continental China, Taiwan, northern Hainan, to northern Vietnam (southernmost distribution) (cf. Table 1; Kamita 1941; Dai et al. 1986; Dai & Yang 1991; Shih 1994; Kosuge et al. 1997; Yoshigou 2001; Shih et al. 2008, 2009)., Published as part of Shih, Hsi-Te, Ng, Peter K. L., Fang, Shou-Hua, Chan, Benny K. K. & Wong, Kingsley J. H., 2010, Diversity and distribution of fiddler crabs (Brachyura: Ocypodidae: Uca) from China, with new records from Hainan Island in the South China Sea, pp. 1-19 in Zootaxa 2640 on page 10, DOI: 10.5281/zenodo.198567, {"references":["Milne Edwards, H. (1852) Observations sur les affinites zoologiques et la classification naturelle des Crustaces. Annales des Sciences naturelles, (3) 18, 109 - 166.","Stimpson, W. (1858) Crustacea Ocypodoidea. Prodromus descriptionis animalium evertebratorum, quae in Expeditione ad Oceanum Pacificum Septentrionalem a Republica Federata missa, Cadwaladaro Ringgold et Johanne Rodgers ducibus, observavit et descripsit, Pars 5. Proceedings of the Academia of Natural Science, Philadelphia, 10, 93 - 110.","Kingsley, J. S. (1880) Carcinological notes, no. II. - Revision of the Gelasimi. Proceedings of the Academy of Natural Sciences of Philadelphia, 1880, 135 - 155.","Cano, G. (1889) Crostacei brachiuri ed anomuri raccolti nel viaggio della \" Vettor Pisani \" intorno al globo. Bollettino della Societa di Naturalisti in Napoli, 3, 79 - 106, 169 - 268.","Shen, C. - J. (1940) The brachyuran fauna of Hong Kong. Journal of the Hong Kong Fisheries Research Station, 1, 211 - 242.","Balss, H. (1922) Ostasiatische Decapoden. IV. Die Brachyrhynchen (Cancridea). Archiv fur Naturgeschichte, Berlin, 88 A (11), 94 - 166.","Gee, N. G. (1926) Tentative list of Chinese decapod Crustacea. Lingnaam Agricultural Review, 3 (2), 156 - 166. [dated 1925]","Kellogg, C. R. (1928) Crustacea of Fukien Province. Lingnan Science Journal, 5 (4), 351 - 356.","Gordon, I. (1931) Brachyura from the coasts of China. Journal of the Linnean Society (Zoology), 38, 525 - 558.","Shen, C. - J. (1936) On a collection of brachyuran Decapoda from Hainan Island with descriptions of three new species.","Jones, D. S. & Morton, B. (1994) The fiddler crabs (Ocypodidae: Uca) of Hong Kong. Asian Marine Biology, 11, 9 - 40.","Yamaguchi, T. (1994) Fiddler crabs of the genus Uca in the collections of three natural history museums in Europe. 1. The specimens held by the Nationaal Natuurhistorisch Museum, Leiden and the Natural History Museum, London. Calanus, 11, 151 - 189.","Cai, L. - Z., Tam, N. F. Y. & Wong, Y. S. (1998) Characteristics of quantitative distribution and species composition of macrozoobenthos in mangrove stands in eastern Hong Kong. Journal of Xiamen University (Natural Science), 37, 115 - 121. [In Chinese]","Ying, X. - P., Zhang, Y. - P., Huang, M. - J. & Yao, Z. - Y. (1999) Floristics trait of intertidal crabs of the islands in the south of Zhe Jiang. Transactions of Oceanology and Limnology, 1999 (4), 31 - 37. [In Chinese]","Xiao, H. - H. & Li, F. - M. (2003) The composition and ecological distribution of intertidal crabs of Naozhou Island in Guangdong. Guizhou Science, 21 (4), 59 - 62. [In Chinese]","Kwok, W. P. W. & Tang, W. S. (2006) Fiddler crabs in Hong Kong - an overview. Hong Kong Biodiversity, 12, 1 - 7.","Yang, S. - L., Chen, H. - L. & Jiang, W. (2008) Brachyura. In: Liu, R. - Y. (Ed.), Checklist of Marine Biota of China Seas. Science Press, Beijing, China, pp. 761 - 810. [In Chinese]","Shen, C. - J. & Dai, A. - Y. (1964) Illustrations of Animals in China (Crustacea Part II). Science Press, Beijing, China. [In Chinese]","Yang, D. - W. & Liu, W. - J. (1996) Crabs animal from the coast of Naozhou Island in Zhanjiang. Journal of Huaihua Teachers College, 15 (6), 187 - 189. [In Chinese]","Crane, J. (1975) Fiddler Crabs of the World (Ocypodidae: Genus Uca). Princeton University Press, Princeton, New Jersey.","Dai, A. - Y., Yang, S. - L., Song, Y. - Z. & Chen, G. - X. (1986) Crabs of the China Seas. China Ocean Press, Beijing, China. [In Chinese]","Dai, A. - Y. & Yang, S. - L. (1991) Crabs of the China Seas. China Ocean Press, Beijing, China.","Lu, L. (1996) Species composition and distribution of intertidal crabs in Xiamen area. Journal of Oceanography in Taiwan Strait, 15 (2), 163 - 169. [In Chinese]","Lai, T. - H. & He, B. - Y. (1998) Studies on the macrobenthos species diversity for Guangxi mangrove areas. Guangxi Sciences, 5 (3), 166 - 172. [In Chinese]","Liang, C. - Y., Zhang, H. - H., Xie, X. - Y. & Zou, F. - S. (2005) Study on biodiversity of mangrove benthos in Leizhou Peninsula. Marine Science, 29 (2), 18 - 25, 31. [In Chinese]","Li, J. - S. (2004) An investigation of the crabs in Mindong tide beach and the initiative of their ecology. Journal of Ningde Teachers College (Natural Science), 16 (1), 21 - 23. [In Chinese]","Chen, Y. - S. (1991) Brachyura (Brachyrhyncha). In: Wei, C. - D. & Chen, Y. - S. (Eds.), Fauna of Zhejiang. Crustacea. Zhejiang Science and Technology Publishing House, Hangzhou, Zhejiang, China, pp. 349 - 387, 402 - 443. [In Chinese]","Naderloo, R., Turkay, M. & Chen, H. - L. (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, 1 - 38.","Kamita, T. (1941) Studies of the Decapod Crustaceans of Chosen. Part I. Crabs. The Fisheries Society of Chosen, Keijo, Korea. [in Japanese]","Shih, H. - T. (1994) Fiddler Crabs. National Museum of Marine Biology / Aquarium, Pingtung, Taiwan. [In Chinese]","Kosuge, T., Wada, K. & Trong, P. D. (1997) Crab distribution in the Cam River estuary, Haiphong, northern Vietnam. In: Anh, P. N., Brands, J. T. & Hong, P. N. (Eds.), National Workshop on the Relationship Between Mangrove Rehabilitation and Coastal Aquaculture in Vietnam. Cres & Actmang, Hanoi, pp. 178 - 184.","Yoshigou, H. (2001) Japanese species of genus Uca (Crustacea: Decapoda: Ocypodidae) and distribution of U. lactea in Hiroshima Prefecture, Japan. Hibakagaku, 200, 1 - 12. [in Japanese]","Shih, H. - T., Lee, J. - H. & Shy, J. - Y. (2008) Three freshwater and coastal crabs (Crustacea: Decapoda: Brachyura) new to Penghu, Taiwan, with notes on their biogeographical implications. Collection and Research, 21, 39 - 44.","Shih, H. - T., Kamrani, E., Davie, P. J. F. & Liu, M. - Y. (2009) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U. lactea species-complex. Hydrobiologia, 635, 373 - 382."]}

Uca annulipes (H. Milne Edwards, 1837) Gelasimus annulipes H. Milne Edwards, 1837: 55; pl. 18, figs. 10���13 [type locality ���Mer des Indes���]. Uca (Celuca) lactea annulipes ��� Crane 1975: 299, pls. 39 A���D, 40 C, D, 45 A, 50 A, figs. 18 A���C, 19 I ���N, 20 D���K, 24 N, O, 27 I, J, 29 D, 32 L, M, 37 L, 41 A, 46 L, 54 I, II, 69 D, 81 O, 83 B [list]; Dai et al. 1986: 426, pl. 59 (3), fig. 236 (2) [Hainan]. Uca (Celuca) annulipes ��� Dai & Yang 1991: 467, pl. 59 (3), fig. 236 (2) [Hainan]. Uca (Paraleptuca) annulipes ���Ng et al. 2008: 241 [list]. Uca annulipes ��� Yang et al. 2008: 807 [Hainan]. Uca (Austruca) annulipes ��� Naderloo et al. 2010: 7, figs. 2 a���h, 3 a���e, 4 b, 12 a, b, c [Hainan]. Material examined. Hainan: 1 3 (16.29 mm), one chela (MBMCAS 166253), Sanya, Hainan, Nov. 1959; 6 3 (10.66���19.81 mm), 1 �� (12.63 mm) (NCHUZOOL 13244), coll. H.-T. Shih & J.-H. Lee, Sanya, 28 Jun. 2004. Distribution. Widely distributed from the tropical West Pacific to the central Indian Ocean (Philippines, Hainan in China, Indochina, Borneo, Malay Peninsula, Sulawesi, Thailand, India). Remarks. In China, this species is only found in southern Hainan in the northern South China Sea (Table 1) and this is probably the northern limit of its range (cf. Shih et al. 2009: fig. 1)., Published as part of Shih, Hsi-Te, Ng, Peter K. L., Fang, Shou-Hua, Chan, Benny K. K. & Wong, Kingsley J. H., 2010, Diversity and distribution of fiddler crabs (Brachyura: Ocypodidae: Uca) from China, with new records from Hainan Island in the South China Sea, pp. 1-19 in Zootaxa 2640 on page 6, DOI: 10.5281/zenodo.198567, {"references":["Milne Edwards, H. (1834 - 1837) Histoire naturelle des Crustaces comprenant l'anatomie, la physiologie et la classification de ces animaux. Paris, Librairie Encyclopedique de Roret. Vol. 1: i - xxxv, 1 - 468. Vol. 2: 1 - 531. Atlas, 1837: 1 - 32, pls. 1 - 42. Vol. 3, 1840: 1 - 638.","Crane, J. (1975) Fiddler Crabs of the World (Ocypodidae: Genus Uca). Princeton University Press, Princeton, New Jersey.","Dai, A. - Y., Yang, S. - L., Song, Y. - Z. & Chen, G. - X. (1986) Crabs of the China Seas. China Ocean Press, Beijing, China. [In Chinese]","Dai, A. - Y. & Yang, S. - L. (1991) Crabs of the China Seas. China Ocean Press, Beijing, China.","Yang, S. - L., Chen, H. - L. & Jiang, W. (2008) Brachyura. In: Liu, R. - Y. (Ed.), Checklist of Marine Biota of China Seas. Science Press, Beijing, China, pp. 761 - 810. [In Chinese]","Naderloo, R., Turkay, M. & Chen, H. - L. (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, 1 - 38.","Shih, H. - T., Kamrani, E., Davie, P. J. F. & Liu, M. - Y. (2009) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U. lactea species-complex. Hydrobiologia, 635, 373 - 382."]}

Uca (Austruca) cryptica sp. nov. (Figs. 5, a���k, 7 b, 8 b) Holotype. 1 male (ZSM), Indonesia, West Flores, mangroves, coll. date and collector unknown. Paratypes. 1 male [(SMF 17167) CL 8.5, CB 15.9 mm], Phillipines, Naawan, ded. K. Sakai; 2 males (NHMW), Indonesia, Celebes, 0 1.1894, leg. Wolf. Diagnosis. Front broad. Anterolateral margin moderately convergent (Fig. 5 a). Major male chela palm with supramarginal groove, partly beset with setae (Fig. 5 e). G 1 with palp, relatively short, hardly reaching to base of horny endpiece (Fig. 5 i). This last one flat, with flanges not inclined, about as broad as long, strongly bent in lateral direction, forming much smaller angle with the stem. Inclination of endpiece is much stronger than in any other species of the lactea -group, Position and displacement of suture as in U. annulipes (Figs. 5 ij). Median plate of the gastric mill with 6 teeth (Fig. 7 b). Description. Carapace smooth. Front broad (about 1 / 7 times as broad as carapace). Venterolateral margin of carapace moderately convergent (Fig. 5 a), distinctly crested; posterolateral margin distinctly crested, extending curvedly backwards, ending in level of central cardiac region; posterolateral striae short. External orbital angle acute, long triangular, directed antero-externally, tip very slightly incurbed frontally. Upper orbital border sinuous, with two crests bordering eyebrow, in its broadest part about 1 / 3 breadth of eyestalk. Lower orbital border (Fig. 5 b) regularly granulate, with granules increasing in size from inner to outer part. Pterygostomian regions beset regularly with feathered bristles, becoming scarce towards lower orbital border (Fig. 5 b). Third maxillipeds with merus 1 / 3 length of ischium. longitudinal broad groove on ischium and merus, close to inner margin, bordered with short setae; long setae along inner margin of ischium, merus. Major chela merus (Fig. 5 c) with scarce granules on anterior upper portion of outer surface; inner dorsal margin with large granules; lower margin denticulate, denticles becoming large dorsally (Fig. 5 d). Carpus definitively longer than broad, outer surface smooth, anterior upper margin with a row of granules, alternating in size; inner surface with scattered granules. Palm (Fig. 5 e) with outer surface minutely granular; lower margin with row of large granules; supramarginal groove adjacent to lower margin, beset with short setae; inner surface (Fig. 5 f) with oblique tuberculate ridge, high, with large granules; proximal ridge at dactylus base parallel to distal one, both strongly tuberculate. Dactylus high in proximal part, higher than fixed finger; both fingers with enlarged proximal tooth; subdistal tooth on fixed finger. Small chela (Fig. 5 g) with smooth ridge on outer surface, extending from lower distal end of palm to proximal two thirds of fixed finger; without teeth on cutting edge of fingers, gap large, broader than adjacent dactylus. Second to fifth pereiopods with merus having convex borders, granular crests on anterior margin (Fig. 5 h); remaining joints smooth. Dactylus moderately curved. Sternum smooth, transverse lines not reaching midline, longitudinal line present in thoracic segments 6-8. Episternites not completely separated from respective sternites. Male abdomen tapering from segments 2 to 5, segment 6 with subparallel margin; length of all segments between 3 and 6 subequal. G 1 with stem (Fig. 5 i) slightly bent in dorso-lateral direction; palp rather short, hardly reaching to base of horny endpiece. This last one about as long as broad, flat, with flanges not inclined, distal border oblique (Fig. 5 j), with terminal opening of sperm channel in midline; dorsal lobe reaching slightly beyond ventral one; whole endpiece bent in lateral direction, thus forming rather small angle with stem, its inclination is much larger than in any other species of the lactea -group. Suture of sperm channel displaced ventrally from base to tip (Fig. 5 k). Median tooth plate of the gastric mill (Fig. 7 b) with 6 teeth; two first ones massive, different in shape; 3-5 decreasing in size distally; third one clearly narrow, large gap between it and second and third teeth; 4 and 5 nearly in the same size, small gap between them; last one shorter, attached to plate dorsally. Lateral tooth plate (Fig. 8 b) with 17 comb-shaped teeth. Distribution. Up to now only known from Indonesia and the Philippines. Remarks. The new species is distinct from any other lactea -group members by the peculiar shape of its G 1, particularly the strongly inclined endpiece. There are other characters that allow its identification. The presence of a supramarginal groove on the major male chela palm leads to the exclusion of all similar species except U. annulipes and U. perplexa. It is distinguishable from these two species by the fact that the new species lacks a tuft of setae on the lower anterior corner of the merus of the large chela, which is present in U. perplexa and U. annulipes. As only the holotype and two paratypes (NHMW) can be compared in this respect, the Philippines paratype lacking the large chela, the identification has always to be determined by examining the G 1. Regarding the median tooth of the gastric mill (Figs. 7 a, b), this species is closely related to U. iranica by having 6 teeth on the median plate, but readily distinguishable from all other members of the group. Considering the general features of the apical part of the G 1 and median tooth plate of the gastric mill, Uca cryptica sp. nov. can be properly attributed to Clade W (this clade already includes U. albimaba, U. annulipes and U. iranica) proposed by Shih et al. (2009). Nothing is known about the biology and ecology of the new species, and very few specimens are known. Etymology. We name it cryptica to point out the fact that it was hidden among numerous museum specimens and still remained undiscovered., Published as part of Naderloo, Reza, T��rkay, 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, pp. 1-38 in Zootaxa 2500 on pages 12-14, DOI: 10.5281/zenodo.195791, {"references":["Shih H. - T, Kamrani E., Davie P. J. F & Liu M. Y. (2009) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U. lactea species-complex. Hydrobiologia, 635, 373 - 382."]}

Uca (Austruca) perplexa (H. Milne Edwards, 1852) (Figs. 16 a���g, 17 a, b, 18 c���f) Gelasimus perplexus H. Milne Edwards 1852: 150. Gelasimus annulipes ��� Kingsley 1880: 148. Uca annulipes var. orientalis ��� Nobili 1901: 13, 14, fig. A. ��� Maccagno 1928: 36, fig. 21. Uca lactea ��� Musgrave 1929: 342, 343 [as lacteae]. ��� Boone 1934: 199, pl. 103. Uca (Celuca) lactea perplexa ��� Crane 1975: 300. figs. 18 D���F, 19 B���H, 20 A���C, 26 D, 31 E, 54 K, 69 C, Pl. 39 E���F. ��� Miyake 1983: 163, pl. 55. fig. 1. Uca (Celuca) lactea lactea ��� T. Sakai 1976: 608. [part.] Uca (Celuca) lactea annulipes ��� T. Sakai 1976: 608. [non Gelasimus annulipes H. Milne Edwards, 1837.] Uca lactea perplexa ��� Shih et al. 1999: 170. Uca perplexa ��� Naiyanetr 1980: 43 (in list). ��� George & Jones 1982: 10 (in key), 83���86, figs. 6, 7b, 54 a���f, 58 e. ��� Ng et al. 2001: 37 (in list). ��� Davie 2002: 362. ��� Naiyanetr 2007: 132 (in list). ��� Shih et al. 2009: 377. Material examined. India: 1 male (UZM), Nicobar Islands, leg. Reinhardt, "Galathea"-Expedition. Thailand: 1 male (UZM), Malay Peninsula, 1915; 1 male (UZM), Koh Chang, stony coast at low water, 0 1.1900, leg. T. Mortensen. Cambodia: 4 males, 1 female (UZM), Lam Ngob, mangrove swamp, 23 ��� 27.12.1899, 1 eg. T. Mortensen. Japan: 1 male (SMF 35021), exact locality unknown, ded. K. Sakai; 1 male (SMF 35022), exact locality unknown, ded. K. Sakai; 1 male, 1 female (SMF 35023), exact locylity unknown, ded. K. Sakai; 3 males (SMF 35024), Ryukyu Islands, Kabira, Ishigaki Island, November 1971, coll. Sakurai, ded. K. Sakai; 2 females (SMF 35018), Amami-Oshima, Tekebu, 24.07. 1966, ded. K. Sakai; 4 males (SMF 35025), Okinawa, ded. K. Sakai; 50 males, 24 females (1 ovig.) (SMF 35012), Ryukyu Archipelago, Yaeyama Islands, Ishigakijima, April 1973, leg. Y. Ono, ded. K. Sakai; 3 males, 3 males (SMF 35019), Ryukyu Archipelago, Yaeyama Islands, Ishigaki-jima, ded. K. Sakai; 15 males, 2 females (SMF 35020), Yaeyama Islands, Ishigaki-jima, Aichi, 12.02. 1995, ded. K. Sakai; 10 males (SMF 17147), Yaeyama Islands, Ishigaki-jima, 1 eg. T. Sakai; 4 males, 3 females (SMF 17146), same locality, 0 7.1971, leg. K. Sakai; 16 males, 10 females (SMF 35014), Yaeyama Islands, Irimote-jima, Ohara, April 1973, Y. Ono, ded. K. Sakai; 4 males (SMF 35015), K. Sakai; 3 males, 1 female (SMF 35016), Iriomote-jima, Ohara, K. Sakai; 3 males, 1 female (SMF 35017), Iriomotejima, Ohara, K. Sakai. Singapore: 4 males (UZM), Putan Sudang, tidal zone, bottom coral reef, 19.05. 1951 ["Galathea"- Expedition. Stat. 337.] Indonesia: 2 males (SMF 5680), Sumatra, Sipora, 1 eg. Maass; 1 male (ZMG 101), Noordwachter Isl. (= Pulo Sebiri) E. of Lampong, 04-05.1885, 1 eg. J. Brock; 1 male (UZM), Padang, 1 eg. Djellerus; 2 males (ZSM), Java, 1929; 3 males (MNHN B 12006), 1 male (MNHN B 12008), 1 male (MNHN B 12009), 1 male (MNHN B 12010), Jakarta, 1 eg. Blecker; 7 males, 1 female (UZM), Tjilatjap; 3 males, 1 female (UZM), Soelyi, mangrove beach, 0 1.04. 1909, "T. Mortensne's Java expedition"; 1 male (ZMG 100), Borneo, Pontianac, 1894, 1eg. Storm; 1 male (MNHN), E. coast, delta of Makaham river, Senipah, 10.1986, 1 eg. Detrieux; 3 males (NHMB 575), Celebes, Kema, 1894, ded. Sarasin; 6 males (ZSM), W-Flores, mangrove beach; 1 male (UZM), Banda Archipelago, Lontor, coast near Lavvande, sand, 0 6.06. 1922, "Danish expedition to Kei Islands 1922 "; 1 male (MNHN B 11882), Aroe islands, 0 6.1959, 1 eg. Lorpuin; 3 males (ZMG 99), Molucca Archipelago, Ambon, 07���09.1885 1 eg. J. Brock; 4 male (SMF 5677), Ternate, 1894, leg. W. K��kenthal; 1 male (NNM 32609), W. New Guinea, Mangroves near Kampong Mandong, between Mokmer and Bosnek, SE Biak, 20.03.1955, 1eg. L. B. Holthuis. Papua New Guinea: 1 male (NHMB 574 d), Bismarck Archipelago, Tatau Island near New Ireland, 1933, ded. A. B��hler; 5 males (ZMH K 5768), New Britain, Gazelle Peninsula, Blanche bay, W. coast of volcanic island, 0 4.08.1908, 1eg. G. Duncker. Australia: 1 male (SMF 17149), Queensland, Bribie Island, N. of Brisbane, W. coast, Banksia beach, mangroves at Dux creek, 31.05. 1980, leg. M. T��rkay; 10 males, 2 females (ZMH), 3 males (SMF 17148), Tannum Sands 25 km from Gladstone, sand mangroves, 28.01. 1976, leg. G. Hartmann & G. Hartmann- Schr��der; 1 male (SMF 17158), Innisfail, 28.04.1957, 1eg. H. Felten; 4 males, 1 female (SMF 17150), Yarrabah NE of Cairns, beach, sand mangroves, 0 6.06. 1980, leg. M. T��rkay; 3 males, 2 females (SMF 17151), Yule point near Port Douglas, N. of Cairns, mangroves, 0 8.06.1980, 1eg. M. T��rkay. Solomon Islands: 2 males (MNHN B 11968). Vanuatu: 2 males (MNHN B 7917), New Hebrides, 0 3.10. 1971, leg. R. Ser��ne. New Caledonia: 6 males (MNHN B 12011); 4 males (MNHN B 12013), 1862; 2 males (MNHN B 12007), 1903, leg. A. Milne Edwards; 1 male (MNHN B 20301), St. 26, 26.09.1978; 1 male (UZM), mouth of Duinbea river, mangrove swamp, 0 4.12. 1928 ["Dana"-Station 3618]; 3 males (MNHN B 20307), 22.09.1978; 22 males (MNHN B 9473), Route de Oucos, 4 km from Noumea, 0 8.08. 1971, leg. R. Ser��ne. Fiji: 1 male (Ex ZMH K 3184); 3 males, 3 females (UZM), Nukumaroreko Isl., Suva Bay, 18 �� 07'S, 178 �� 24 'E, below high water mark on dry sand, 1 eg. T. Wolf. Tonga: 1 male (ZMH K 2914); 2 males (UZM), Nukualofa, intertidal, 0 1.1956, leg. Mischele. Samoa: 1 male (ZMH K 2909); 1 male (ZMH K 2915); 4 males (SMF 5675), 1904, leg. Hauck; 1 male (SMF 17153), Upolu, vend. Mus. Goddefroy. Pacific Ocean: 1 male (UZM), Mus. Goddefroy, ded. Wroblewsky; 1 male (ZMK Cr. 1528), 1876, leg. G. Jantzen. Wrong or doubtful localities: 1 male (ZNIH K 2938), "South Atlantic". Unknown locality: 4 males (ZMH K 6748), vend. J. Umlauff, 0 4.03. 1914. Diagnosis. Front broad. Anterolateral borders moderately convergent. Major male chela with palm bearing a supramarginal groove adjacent to lower border, beset with short setae (Figs. 16 a, b). G 1 (Fig. 16 d) with palp long, clearly reaching beyond base of horny endpiece; this last one has inclined flanges, strongly folded towards dorsal direction in comparison to the stem (Fig. 16 f); terminal opening of sperm channel at base of shallow emargination of the distal margin (Fig. 16 e). Genital opening of female (Fig. 16 g) and thus also direction of opercle edge oblique in respect to median line of sternum. Distribution. Eastern Indian Ocean (from the Nicobar Is. eastward), western and central Pacific Ocean as far north as the Ryukyu Islands to Amami-Oshima. In Australia, the species is distributed throughout the east coast reaching its southern limit around Trial Bay in southern New South Wales. Remarks. Crane (1975) reported one or two intermediate individuals from north-central Philippines, and believed that these emerged from hybridization between U. perplexa and U. annulipes. She referred mainly to the intermediate form of the G 1 of these specimens, and mentioned that interbreeding occurs between these two species in Sunda Shelf. Subsequently, Barnwell (1980) examined specimens of U. perplexa and U. annulipes from Jakarta, Indonesia, and found three different forms contributing to U. perplexa, U. annulipes and an intermediate form. He believed that all three are distinct species, but this intermediate form never has been described as a new species. We examined numerous specimens of U. perplexa from different locations within its distribution as far as South Australia and Japan, including areas where it was thought to occur sympatrically with U. annulipes. Uca perplexa deserves specific rank, as G 1 and the female genital opening are in all cases different from U. annulipes and allies, also from U. lactea and U. mjoebergi. The strong torsion of the endpiece, together with its prominent emargination is especially distinctive in Uca perplexa. The corresponding shape of the female genital opening also allows the recognition of females. The median tooth plate of the gastric mill is a further character that is used to separate this species from its congeners. We, therefore, do not agree with Crane���s (1975) hypothesis that interbreeding occurs between U. perplexa, and U. annulipes. However, U. perplexa is morphologically more similar to U. lactea than to U. annulipes. They share similarities in the G 1, major chela of the male and the morphology of the median tooth plate of the gastric mill. Genetic results of Shih et al. (2009) support this idea by grouping U. perplexa with U. lactea in the same clade (Clade E by Shih et al. 2009). They also treated U. perplexa as a good species as did George & Jones (1982)., Published as part of Naderloo, Reza, T��rkay, 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, pp. 1-38 in Zootaxa 2500 on pages 24-30, DOI: 10.5281/zenodo.195791, {"references":["Milne Edwards H. (1852) Observation sur les affinites zoologiques et la classification naturelle des crustaces. Annales des Sciences Naturelles (Zoologie), (3) 18, 109 - 166.","Kingsley J. S. (1880) Carcinological notes. Proceedings of the Academy of Natural Sciences of Philadelphia. For 1879: 34 - 37; ii. Revision of the Gelasimi, 135 - 152.","Nobili G. (1901) Note intorno ad una collezione di crostacei di Sarawak (Borneo). Bolletino di Zoologia ed Anatomia comparata. Della R. Universita di Torino, 16 (397), 1 - 14.","Maccagno T. (1928) Crostacei decapodi. Le specie del genre Uca Leach conservate nel Regio Museo Zoologico di Torino. Bolletino dei Musei di Zoologia e di Anatomia Comparata della R. Universita di Torino, 41 (11), 1 - 52.","Musgrave A. (1929) Life in a mangrove swamp. Australian Museum Magazine, 3 (10), 341 - 347.","Boone L. (1934) Scientific Results of the World Cruise of the Yacht \" Alva \", 1931, William K. Vanderbilt, Commanding. Crustacea: Stomatopoda and Brachyura. Bulletin of the Vanderbilt Marine Museum, 5, 210 pp. Borradaile L. A. (1907) Land crustaceans. In: Gardiner, J. S. [Ed.], Fauna and Geography of the Maldive and Laccadive Archipelagoes. Being the Account of the Work carried out on and of the Collections made by an Expedition during the years 1899 and 1900. Volume 1, Cambridge University Press, Cambridge, pp. 65 - 100.","Crane J. (1975) Fiddler Crabs of the World. Ocypodidae: Genus Uca. Princeton University Press, Princeton, New Jersey, i - xxiii + 736 pp.","Miyake S. (1983) Japanese crustacean decapods and stomatopods in color. 1. Macrura, Anomura and Stomatopoda. Osaka: Hoikusha. 261 pp.","Sakai, T. (1976) Crabs of Japan and the Adjacent Seas. Kodansha, Tokyo, vol. 1 [English text], xxix + 773 pp., figs. 1 - 379, maps 1 - 3; vol. 2 [Japanese text], 461 pp., figs. 1, 2; vol. 3 [plates], 16 pp. + pls. 1 - 251.","Milne Edwards H. (1837) Histoire naturelle des Crustaces, comprenant l'Anatomie, la Physiologie et la classification de ces animaux. ii. Paris, 1837, 531 pp.","Shih H. - T, Mok H. - K, Chang H. - W & Lee S. - C. (1999) Morphology of Uca formosensis Rathbun, 1921 (Crustacea: Decapoda: Ocypodidae), an endemic fiddler crab from Taiwan, with notes on its ecology. Zoological Studies, 38 (2), 164 - 177.","Naiyanetr P. (1980) Crustacean Fauna of Thailand (Decapoda and Stomatopoda). Department of Biology, Faculty of Science, Chulalongkorn University of Bangkok, Thailand, 73 pp.","George R. W. & Jones D. S. (1982) A revision of the fiddler crabs of Australia (Ocypodidae: Uca). Record of the Western Australian Museum, Suppl. 14, 99 pp.","Naiyanetr P. (2007) Checklist of crustacean fauna in Thailand (Decapoda, Stomatopoda, Anostraca, Myodocopa and Isopoda). Open Biodiversity Series, 19, 197 pp.","Shih H. - T, Kamrani E., Davie P. J. F & Liu M. Y. (2009) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U. lactea species-complex. Hydrobiologia, 635, 373 - 382.","Barnwell F. H. (1980) Taxonomic differentiation within the lactea - complex of fiddler crabs in the Sunda shelf region. American Zoologist, 20, 957 [abstract]."]}

Uca (Austruca) iranica Pretzmann, 1971 (Figs. 6, c���f, 7 a, 8 a, 9 a���k) Uca annulipes ��� Stephensen 1945: 189. ��� Pretzmann 1971: 481 [not Gelasimus annulipes H. Milne Edwards, 1837]. ��� Al-Ghais & Cooper 1996: 419 ���421, figs. 7���8. ��� Tirmizi & Ghani 1996: 105 ���107 (not fig. 40). ��� Hornby 1997: 15 (prt.?). [not Gelasimus annulipes H. Milne Edwards, 1837] Uca annulipes iranica Pretzmann 1971: 481, P 1. 5, figs. 11���12. ��� Apel & T��rkay 1999: 133 ���134. ��� Apel 2001: 114. Uca (Celuca) lactea annulipes ��� Crane 1975: 299, 301, 611. [part: Persian Gulf���s material]. ��� Collins et al. 1984: 318, 326 ��� 327. ��� Jones 1986: 157, pl. 42. Austruca lactea annulipes ��� Clayton 1986: 86 ���88, fig. 3. Uca iranica ��� Shih et al. 2009: 377. Holotype. 1 male (NHMW 3788), Iran, Bandar-Abbas, 1970, leg. Pretzmann & Bilek. Other material. Persian Gulf: Iran: 3 males (NHMW), Bandar-Abbas, intertidal beach, 1970, leg. Pretzmann & Bilek; 24 males, 4 females, 3 juv. (SMF 34650), Qeshm Island, N. coast, 3 km W of Kuweii, 26 �� 57 'N, 56 �� 00'E, muddy flat sometimes mixed with shells, among sea grass, 0 6.05. 2008, leg. R. Naderloo & A. Kazemi; 4 males, 2 females, 12 juv. (SMF 34651), Qeshm Island, S. coast, 26 �� 43 ��N, 55 �� 49 ��E, sandy flat, 0 9.01. 2008, R. Naderloo & M. T��rkay; 21 males, 5 females (SMF 34652) Qeshm Island, Dargahan, 26 �� 58 ��N, 56 �� 0 4 ��E, sandy-muddy, with planted mangroves, 13.01. 2008, R. Naderloo & M. T��rkay; 6 males, 2 females (SMF 34653), Qeshm Island, S. coast, 26 �� 48 ��N, 56 �� 0 5 ��E, sandy, muddy-sand substrate with dead plantedmangroves, 11.01. 2008. R. Naderloo & M. T��rkay; 9 males, 6 females (SMF 34654), Qeshm Island, N. Coast, Holor village, 26 �� 38 ��N, 56 �� 0 4 ��E, sandy substrate, planted mangroves, 13.02. 2006, R. Naderloo; 6 males, 9 females (ZUTC Brach 1246), Qeshm Island, Tula, 26 �� 59 ��N, 56 �� 12 ��E, muddy-sand, 13.01. 2008, R. Naderloo & M. T��rkay; 9 males, 2 females (ZUTC Brach 1247), Bushehr, Shah Abdollah, 30 �� 0 6 ��N, 49 �� 46 ��E, sandymud flat, 14.04. 2006, R. Naderloo & A. Kazemi; 2 males (ZUTC Brach 1248), Bushehr, Delvar, Park Saheli, 28 �� 45 ��N, 51 �� 20 ��E, sandy substrate along tidal water channel, 10.04. 2006, R. Naderloo & A. Kazemi; 5 males (ZUTC Brach 1249), Bandar-Emam, Park Saheli, 30 �� 28 ��N, 49 �� 0 4 ��E, muddy substrate, 21.05. 2008, R. Naderloo & A. Kazemi; 2 males, 1 female (ZUTC Brach 12450), Bandar-Abbas, behind Airport, 27 �� 47 ��N, 56 �� 24 ��E, tidal channel with planted mangroves, sandy-mud substrate, 0 7.01. 2008, R. Naderloo & A. Sari. Kuwait: 1 male (NHM 1988: 54), 1981, leg. D.A. Jones; 2 males (SMF 17154), 1 eg. A. Seilacher; 7 males, 1 female (NHM 1988: 53), 29 �� 35 'N, 48 �� 09'E, leg. D.J. Crisp; 3 males, l female (NHM 1978: 197), Kuwait Bay, Al Jurthamah, intertidal sand, 0 4.04. 1978, leg. D. Clayton & T. Vaughan; 1 male, 1 female (NHM 1988: 52), Kuwait Bay, Al Akkas, 29 �� 22 'N, 47 �� 55 'E, leg. D.J. Crisp. UAE: 3 males (SMF 26026), Ras al Khaimah, mangroves, N. Rams, 25 �� 50 'N, 55 �� 00'E, sandy-muddy substrate, high intertidal, 10.07. 1995, leg. M. Apel; 7 males, 2 females (SMF 26027), Ras al Khaimah, inner side of S. Rams, 25 �� 50 'N, 55 �� 00'E, sandy substrate amongst Salicornia sp., 11.07. 1995, leg. M. Apel; 7 males, 1 female (SMF 26024), Ras al Khaimah, Khor, 25 �� 46 'N, 55 �� 58 'E, E. Police club, mangroves, 10.07. 1995, leg. M. Apel; 3 males, 1 female (SMF 26030), Um al Qaiwain, Khor al Beidah, mangrove region, 25 �� 35 'N 55 �� 35 'E, 0���1 m, several small islands, 0 9.07. 1995, leg. M. Apel. Gulf of Oman: Iran: 4 males (SMF 34655), Baluchestan, Guader Bay, mangroves, 25 �� 0 8 ��N, 61 �� 27 ��E, sandy-muddy substrate, 18.11. 2005, R. Naderloo & A. Kazemi; 4 males, 2 females (SMF 34656), 30 km E. Jask, Khor Khalasi, 25 �� 35 ��N, 58 �� 0 2 ��E, seawards fringe of mangroves, muddy-sand substrate, 20.11. 2005, R. Naderloo & A. Kazemi. UAE: 11 males (SMF 26019), Fujairah, mangroves of khor Kalba, 25 �� 01'N, 51 �� 21 'E, sandy-mud substrate, 0 1.07. 1995, leg. M. Apel; 3 males, 4 females (SMF 26023), Fujairah, mangroves of khor Kalba, 25 �� 01'N, 56 �� 21 'E, sandy-mud substrate, 0 1.07. 1995, leg. M. Apel; 3 females (SMF 26022), Fujairah, mangroves of khor Kalba, 25 �� 01'N, 56 �� 21 'E, sandy-mud substrate, 0 1.07. 1995, leg. M. Apel. Diagnosis. Front broad. Anterolateral borders of carapace weakly arched, nearly straight. Major male chela palm without supramarginal groove adjacent to lower border, but shallow depression near base of fixed finger. G 1 and female genital opening as in U. annulipes. Redescription. Front broad, about 1 / 6.5 times as broad as carapace. Venterolateral margin of carapace (Figs. 6 d, 9 a) very slightly convergent, nearly straight, distinctly crested; posterolateral margin crested, extending curvedly backwards, ending about in level of middle cardiac region; posterolateral striae short, shortly behind posterolateral margin, slightly lower that latter. Exorbital angle triangular, directed forward; upper orbital margin sinuous, with two crests (lower one faintly crested), bordering narrow eyebrow, getting strongly narrower outwardly, inner part less than 1 / 2 breadth of eyestalk in adjacent region; lower orbital margin regularly granulate, granules becoming larger on outer one third portion (Fig. 9 b). Pterygostomian region with regular feather-shaped bristle near third maxilliped, becoming glabrous towards lower orbital margin, one longitudinal line of bristle on low-laid antero-inner portion. Third maxilliped with merus about 1 / 3 length of ischium; longitudinal broad groove on ischium, merus, close to inner margin, bordered with short setae; long setae on inner margin of ischium, merus. Major chela merus (Fig. 9 d) with transverse ridges on upper margin, finely granulate; inner upper margin distally elevated, granulate; lower margin denticulate, denticles becoming larger distally (Fig. 9 c). Carpus about 1.5 times as long as broad, inner upper margin denticulate, 3, 4 proximal denticles remarkably large; outer surface, inner surface and lower margin smooth, glabrous. Outer surface of palm (Fig. 9 e) smooth, without supramarginal groove, with shallow depression near base of fixed finger; lower margin weakly granulate, with low granules, upper margin granulate, proximal portion with dense short setae at inner surface; inner surface (Fig. 9 f) with high, oblique and tuberculate ridge, granules large, becoming smaller distally; proximal ridge at dactylus base with large granules, distal ridge parallel to proximal one, with granules low, relatively small. Dactylus more than 2 times as long as palm, slightly wider than fixed finger, with subproximal tooth, median teeth nearly in middle; fixed finger with large median tooth on proximal third, located between proximal and middle teeth of dactylus when fingers are closed, subdistal tooth small. Small chela (Fig. 9 g) with smooth ridge on outer surface, extending from lower distal end of palm to proximal two third of fixed finger; cutting edge without tooth, dactylus slightly wider than gap between fingers, gap getting narrower distally, fixed finger slightly higher than dactylus proximally. Merus of walking legs with finely granulated transverse ridges on anterior margin, posterior margin serrated on proximal half; propodus 1.2 times as long as carpus, propodus of last leg slightly shorter than carpus (Fig. 9 h); dactylus conical, as long as propodus; row of brown bristles on anterior and posterior surface of propodus and dactylus, those of dactylus longer. Male abdomen tapering from segments 2 to 5; segment 6 with lateral margin nearly straight; length of segments 3���5 subequal; segments 5 and 6 of nearly equal length. G 1 (Fig. 9 i) with stem bent in posterolateral direction, palp not reaching to base of horny endpiece; latter slightly longer than wide, distal margin convex, dorsal lobe large (Fig. 9 k), reaching slightly beyond ventral lobe; terminal opening (Fig. 9 j) of sperm channel subdistdal ventrally, in midline of endpiece lobes; feathershaped setae along lateral margin, distally longer, scarce short setae on palp. Genital opening of female as in U. albimana Median tooth plate of the gastric mill (Fig. 7 a) with 6 teeth, with relatively large gaps between them, two first ones massive, different in shape, 3���5 nearly of same size, 6 shorter; lateral tooth plate (Fig. 8 a) with 18 comb-shaped teeth. Distribution. Persian Gulf and Gulf of Oman. Remarks. Pretzmann (1971) originally described the species as a subspecies of U. annulipes from the Persian Gulf. He recorded both U. annulipes and U. iranica from the same locality in Bandar-Abbas. The main characteristics given by Pretzmann (1971) to distinguish the two forms were mainly limited to the morphology of the large chelae. Apparently he assigned specimens with chela without large teeth on the cutting edge of the fingers (mainly regenerated form) to his subspecies U. annulipes iranica, but in the material examined here both species of U. iranica and U. annulipes have broadly the same type of chela. Such differences in the major chelae, which in realty refer to the peculiarities of regenerated pincers, are described also by Yamaguchi (1973) for Uca lactea. We also include in U. iranica the specimens from Bandar-Abbas assigned to U. annulipes proper by Pretzmann (1971). Crane (1975) did not mention or discuss the species. As stated under the remarks for U. annulipes, all examined specimens clearly differ from U. annulipes and U. albimana by the shape of the anterolateral borders. Regarding the median tooth of the gastric mill, this species is clearly different from the two former, but is similar to U. cryptica sp. nov. by having 6 teeth on the median tooth of the gastric mill (Figs. 7 a, b). In contrast, the lateral tooth of the gastric mill of U. iranica is different from U. cryptica sp. nov. and allied to U. albimana and U. annulipes, as U. iranica has a narrow lateral tooth plate with about 18 comb-shaped teeth., Published as part of Naderloo, Reza, T��rkay, 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, pp. 1-38 in Zootaxa 2500 on pages 14-19, DOI: 10.5281/zenodo.195791, {"references":["Pretzmann G. (1971) Ergebnisse einiger Sammelreisen nach Vorderasien. 2. Teil: Marine Brachyura. Annalen des Naturhistorischen Museums Wien, 75, 477 - 487.","Milne Edwards H. (1837) Histoire naturelle des Crustaces, comprenant l'Anatomie, la Physiologie et la classification de ces animaux. ii. Paris, 1837, 531 pp.","Al-Ghais S. M. & Cooper, R. T. (1996) Brachyura (Grapsidae, Ocypodidae, Portunidae, Xanthidae and Leucosiidae) of Umm Al Quwain mangal, United Arab Emirates. Tropical Zoology, 9, 409 - 430.","Tirmizi N. M. & Ghani N. (1996) Marine Fauna of Pakistan: 5: Crustacea: Brachyura, Brachyrhyncha, Part 1: Xanthidae, Goneplacidae, Pinnotheridae, Ocypodidae, Grapsidae. Center of Excellence, University of Karachi, 1 - 188.","Hornby R. (1997) A survey of the habitats, invertebrate fauna and environmental sensitivity of the mainland coast of the UAE, with information on status and distribution of Crustaceans. TRIBULUS, Bulletin of the Emirates Natural History Group, 7 (2), 11 - 17.","Apel M. & Turkay M. (1999) Taxonomic composition, distribution and zoogeographic relationships of the grapsid and ocypodid crab fauna of intertidal soft bottoms in the Arabian Gulf. Estuarine Coastal and Shelf Science, 49 (suppl. A), 131 - 142.","Apel M. (2001) Taxonomie und Zoogeographie der Brachyura, Paguridea und Porcellanidae (Crustacea: Decapoda) des Persisch-Arabischen Golfes. Unpublished Ph. D. Thesis, Johann Wolfgang Goethe-Universitat, Frankfurt am Main, 268 pp.","Crane J. (1975) Fiddler Crabs of the World. Ocypodidae: Genus Uca. Princeton University Press, Princeton, New Jersey, i - xxiii + 736 pp.","Jones D. A. (1986) A Field Guide to the Sea Shores of Kuwait and the Arabian Gulf. University of Kuwait Blandford Press, Poole, Kuwait (1986), 192 pp.","Clayton D. A (1986) Ecology of mudflats with particular reference to those of the northern Arabian Gulf. In: Halwagy, R., Clayton, D. & Behbehani, M. (Eds.), Marine Environment and Pollution. Proceedings of the 1 st Arabian Gulf Conference on Environment and Pollution, Kuwait, 7 - 9 February 1982. University of Kuwait, pp. 83 - 96.","Shih H. - T, Kamrani E., Davie P. J. F & Liu M. Y. (2009) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U. lactea species-complex. Hydrobiologia, 635, 373 - 382.","Yamaguchi T. (1973) Asymmetry and dimorphism of chelipeds in the fiddler crab, Uca lactea De Haan. Zoological Magazine (Tokyo), 82 (3), 154 158."]}

Uca (Austruca) annulipes (H. Milne Edwards, 1837) (Figs. 2 a���h, 3 a���e, 4 b, 12 a, 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; 1887 a: 118; 1887 b, 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 1894 a: 758, 759; 1894 b: 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. 1 a. ��� T. Sakai 1940: 32. ��� Chhapgar 1957: 508 ���509, pl. 13, figs. i���o. Gelasimus annulipes var. lacteus ��� Ortmann 1894 a: 759. Uca annulipes ��� Nobili 1899 a: 274; 1899 b: 518; 1903: 20. ��� Doflein 1899: 193. ��� Crosnier 1965: 117, figs. 204, 206, 207, 212, 213. ��� Lanchester 1900 a: 754; 1900 b: 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, 19 e. ��� Tweedie 1950: 356. ��� Sankarankutty 1961: 113. ��� Forest & Guinot 1961: 141. ��� Claude 1964: 11. ��� Pillay & Nair 1971 a: 153; 1971 b: 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. 18 A���C, 19 I ���N, 20 D���K, 24. ��� Frith 1977: 6, 13 (in list). ��� Vannini & Valmori 1981: 208 ���210, figs. 5 c 1 ���4, 6c. Lectotype. 1 male (MNHN B 11854),"Mer des Indes", leg. Reynaud. Paralectotype. 1 male (MNHN B 11884), data as lectotype. Other material. Mozambique: 1 male (ZMH K 2913); 2 males (ZMH K 11921), 03.08.1927; 1 male (SMF 17138), Beira, 03.09.1962; 1 male, 1 female (ZMH K 12863), E. Africa (possibly Mozambique). Tanzania: 10 males (NHMB 576 c), leg. La Roche, 1910; 4 males (SMF 5682), Dar Es Salam, 1899, 1eg. F. Winter; 8 males (ZSM), 0 9.1890, 1 eg. M��ller; 3 males, 1 female (ZMH K 29767), 10.11.1967; 1 male (ZMH K 2916), Bagamoyo, Lagoon, N. of the city, 26.06. 1888, leg. F. Stuhlmann; 1 male (ZMH K 3362), Kingani, near to ferry, mangroves, burrowing in muddy banks, 29.06. 1888, leg. F. Stuhlmann; 9 males, 1 female (ZMH K 2916), Kingani, near ferry to Bagamoyo, flooding area, mud, 0 8.07. 1888, leg. F. Stuhlmann; 3 males (ZMH Ex K 2993), Zanzibar; 31 males (ZMH K 2898); 5 males (ZMH K 2911); 5 males (ZSM); 8 males, 6 females (ZMH K 2919), Lagoon, 20.04.1888, 1eg. F. Stuhlmann. Kenya: 71 males, 2 females (ZSM), Mombasa, leg. Wache; 1 male (ZMH K 27111), 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, 1 eg. M. Grasshoff; 5 males, 2 females (MNHN B 11873), 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. M��ller. 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 B 11880); 4 males (MNHN 11889); 1 male (MNHN B 11892); 1 male, 1 female (MNHN B 11894); 1 male (MNHN B 11896); 20 females (MNHN B 11904); 4 males (MNHN B 11909); 1 male (MNHN B 11910); 1 male, 1 female (MNHN B 11913); 11 males (MNHN B 11900), 1 eg. Waterlot; 2 males (MNHN B 11899), leg. G. Petit; 11 males, 3 females (SMF 5853), 1880, leg. Stumpf & Ebenau; 1 male (SMF 568 l), 1881, leg. A. Stumpf; 3 males, 1 female (MNHN B 11890), Nossi B��; 1 male (MNHN B 11879), 6 males (MNHN B 11880). 1 male (UZM), leg. A Crosnier; 1 male (MNHN B 2024 l), 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 B 11893), Tamatave; 2 males, 2 females (MNHN 11898), Majunga Bay, 1926; 13.11. 1962, leg. Arnoult; 2 males (MNHN B 11902), Marondava, leg. Fauliat; 4 males, 5 females (MNHN B 11907), Ampalaza Bay, 1 eg. G. Petit; 1 male (MNHN B 11906), 2 males (MNHN B 11908), Analalava, 1903, 1eg. E. HANTZ; 2 males (1 juv) (MNHN B 12085), NW coast, Irohono, mangroves, 21.12. 1965, leg. Betsch. Seychelles: 5 males (SMF 6692), leg. R. Ser��ne; 14 males, 9 females (SMF 12944), Praslin, freshwater creek about 300 m. from sea, 16.05.1979, 1eg. M. Ackermann; 4 males (ZMH K 32308), Mah��, 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 B 11875), 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 K 2977); 5 males (ZMH K 26900); 27 males, 2 females (ZMH K 26913); 3 males (SMF 2343), 1 male (ZSM), Bombay, Bandra, shore, 1 eg. R. Altevogt; 6 males (ZMH K 26870), Alibag S. of Bombay, 28.11. 1955, leg. G.A. Von Maydell; 11 males (ZMH Ex 27091), 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, 1 eg. Berg & Mathiassen; 1 male (UZM), Vellore, 1 eg. Loeventhal; 1 male (MNHN B 11878), Karikal, intertidal, 20.12.1940, 1eg. K. Lindberg; 5 males (UZM), Trankebar, "Galathea"-Expedition; 1 female (MNHN B 11853), Vellar estuary near Porto Novo, 14.05. 1980, leg. P. Noel; 1 female (MNHN B 11881), Pondichery; 17 males, 4 females (MNHN B 11888); 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. K��nig; 4 males (ZMH K 30309), Bentota, river estuary; 2 males (ZMH K 2912), 0 3.08.1909, 1eg. G. Duncker; 5 males (SMF 5692), Weligama-Bay, 0 4.02. 1912, leg. O. L��w-Beer; 7 males (NHMB 576 a), Trincomalee, 1886���1893, leg. Sarasin; 6 males (UZM), shore, 0 3.1889, 1 eg. 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 [5 th 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. Ser��ne; 6 males (ZMH K 3073), 3 males (ZMH K 3204), 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 576 b), 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, 1 eg. Wessel; 1 male (ZMH K 3205), W. Africa, 1911, leg. W. Michaelsen. Unknown locality: 25 males, 1 female (MNHN B 11865); 2 males (1 Chelip of male) (MNHN B 11877); 42 males, 3 females (MNHN B 11886); 1 male (MNHN B 11895); 2 males (MNHN B 11901); 6 males (MNHN B 11903); 1 male (MNHN B 11905); 1 male (MNHN B 11911); 1 male (MNHN B 11937); 4 males (UZM); 1 male (ZMH Ex K 6748); 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. G 1 (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 G 1 s are within the range of variability of U. annulipes (see Figs. 3 b���e, or Crane 1975: fig. 65 D). 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, 7 a). 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 G 1 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., Published as part of Naderloo, Reza, T��rkay, 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, pp. 1-38 in Zootaxa 2500 on pages 7-11, DOI: 10.5281/zenodo.195791, {"references":["Milne Edwards H. (1837) Histoire naturelle des Crustaces, comprenant l'Anatomie, la Physiologie et la classification de ces animaux. ii. Paris, 1837, 531 pp.","Krauss F. (1843) Die Sudafrikanischen Crustaceen. Eine Zusammenstellung aller bekannten, Malacostraca. Bemerkungen uber deren Lebensweise und geographische Verbreitung, nebst Beschreibung und Abbildung mehrerer neuen Arten. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart, 68 pp.","Alcock A. (1900) Material for a carcinological fauna of India. No. 6: The Brachyura Catometopa, or Grapsoidea. Journal of the Asiatic Society of Bengal, 69 (2), 279 - 456.","White A. (1847) List of the specimens of Crustacea in the Collection of the British Museum. London, I - VIII + 143 pp.","Dana J. D. (1852) Crustacea, Part i. In: United States Exploring Expedition during the years 1838 to 1842, under command of Charles Wilkes, U. S. N. C. Sherman, Philadelphia, 13 (1), i - viii + 685 pp.","Milne Edwards H. (1852) Observation sur les affinites zoologiques et la classification naturelle des crustaces. Annales des Sciences Naturelles (Zoologie), (3) 18, 109 - 166.","Heller C. (1865) Crustaceen. In: Reise der osterreichischen Fregatte Novara \" um die Erde, in den Jahren 1857, 1858, 1859 unter den Befehlen des Commodore B. Von Wullerstrof-Urbair. Zoologischer Theil, 2 (3), 280 pp, pls. 1 - 25.","Hilgendorf F. (1869) Crustaceen. In: Baron Carl Claus Von Der Decken's Reisen in Ost-Afrika. Dritter Band: Wissenschaftliche Ergebnisse. Erste Abtheilung: Saugethiere, Vogel, Amphibien, Crustaceen, Mollusken und Echiodermen. C. F. Winter'sche Verlagshandlung, Leipzig & Heidelberg, 3 (1), 67 - 116, pls. 1 - 6.","Hoffmann C. K. (1874) Crustaces et Echinodermes de Madagascar et de l'Ile de la Reunion. In: Pollen, F. P. L. & Van Dam, D. C., Recherches sur la faune de Madagascar et ses dependances, 5 (2), 1 - 58, pls. i - x.","Miers E. J. (1879) On a collection of Crustacea made by Capt. H. C. St. John, R. N., in the Korean and Japanese seas. Part i. Podophthalmia. Proceedings of the Zoological Society of London, 1879, 17 - 61, 3 pls.","Kingsley J. S. (1880) Carcinological notes. Proceedings of the Academy of Natural Sciences of Philadelphia. For 1879: 34 - 37; ii. Revision of the Gelasimi, 135 - 152.","Richters F. (1880) Decapoda. In: Mobius, K. (ed.) Beitrage zur Meeresfauna der Insel Mauritius und der Seychellen bearbeitet von K. Mobius, F. Richters und E. von Martens nach Sammlungen, angelegt auf einer Reise nach Mauritius von K. Mobius. Berlin: Verlag der Gutmannischen Buchhandlung, 139 - 178.","Man J. G. De (1880) On some species of Gelasimus and Macrophthalmus. Notes from the Leyden Museum, 2, 67 - 72.","Lenz H. & Richters F. (1881) Beitrag zur Crustaceenfauna von Madagascar. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 12, 421 - 428.","Miers E. J. (1886) Report on the Brachyura collected by H. M. S. Challenger during the years 1873 - 1876. In: Murray, J. (ed.) Zoology. Report on the Scientific Results of the Voyage of H. M. S. Challenger During the Years 1873 - 76, vol. 17, 362 pp.","Alcock A. (1892) On the habits of Gelasimus annulipes. Annals and Magazine of Natural History, 6 (10), 415 - 416.","Henderson J. R. (1893) A contribution to Indian carcinology. Transactions of the Linnean Society London, 2 (10), 325 - 458, pls. 36 - 40.","Ortmann A. E. (1894 a) Die Decapoden-Krebse des Strassburger Museums. viii. Theil. Abteilung: Brachyura (Brachyura genuina Boas) iii. Unterabteilung: Cancroidea, 2. Section: Cancrinea, 2. Gruppe: Catometopa. Zoologische Jahrbucher, Abtheilung fur Systematik, Geographie und Biologie der Thiere, 7, 683 - 772, pl. 23.","Alcock A. & Anderson A. R. S. (1894) List of the shore and shallow water Brachyura collected during the season 1893 - 94. Natural history notes from H. M. Indian Marine survey steamer \" Investigator \". Ser. ii, No. 17. Journal of the Asiatic Society of Bengal, 63, (4), 197 - 209.","Zehnter L. (1894) Crustaces de 1 ' Archipel malais-voyage de M. M. Bedot en Ch. Pichet dans 1 ' Archipel malais. Revue suisse de Zoologie, 2 (1), 135 - 214.","Lenz H. (1905) Ostafrikanische Dekapoden und Stomatopoden. Gesammelt von Herrn Prof. Dr. A. Voeltzkow. In: Voeltzkow, A.: Wissenschaftliche Ergebnisse der Reisen in Madagaskar und Ostafrika in den Jahren 1889 - 95. Abhandlungen der Senckenbergischen naturforschenden Gesellschaft, 27 (4), 341 - 392.","Sewell R. B. (1913) Notes on the biological work of the R. I. M. S. S. \" Investigator \" during survey seasons 1910 - 1911 and 1911 - 1912. Journal of the Asiatic Society of Bengal, 9, 329 - 390.","Bouvier E. L. (1915) Decapodes marcheurs (Reptantia) et stomatopodes recueillis a l'ile Maurice par M. Paul Carie. Bulletin Scientifique de la France et de la Belgique, 48 (3), 178 - 318.","Kemp S. (1915) Fauna of the Chilka Lake No. 3. Crustacea Decapoda. Memoirs of the Indian Museum, 5, 199 - 325.","Gordon I. (1934) Resultats scientifiques du voyage aux Indes Orientales Neerlandaises de LL. AA. RR. le Prince et la Princesse Leopold de Belgique. Crustacea Brachyura. Memoires du Musee Royal D'Histoire Naturelle de Belgique, Hors Serie 3 (15), 3 - 78.","Tweedie M. W. F. (1937) On the crabs of the family Ocypodidae in the collection of the Raffles Museum. Bulletin of the Raffles Museum, 13, 140 - 170.","Sakai T. (1940) Biogeographic review on the distribution of crabs in Japanese waters. Records of Oceanographic Works in Japan, 11 (1), 27 - 63.","Chhapgar B. F. (1957) On the marine crabs (Decapoda, Brachyura) of Bombay State. Part ii. Journal of the Bombay Natural History Society, 54, 503 - 549.","Nobili G. (1899 a) Contribuzioni alla conscenza della Fauna carcinologia della Papuasia, delle Malucche e dell'Austrialia. Annali del Museo civico di Storia naturale Genova, 20, 230 - 282.","Doflein F. (1899) Amerikanische Dekapoden der k. bayerischen Staatssammlungen. Sitzungsber. Koniglich Bayerischen Akademie der Wissenschaften, 29, 177 - 95.","Crosnier A. (1965) Crustaces Decapodes, Grapsidae et Ocypodidae. Faune de Madagascar, 18, pp 1 - 143.","Lanchester W. F. (1900 a) On a collection of crustaceans made at Singapore and Malacca. Part 1. Crustacea Brachyura. Proceedings of the Zoological Society of London, 3, 719 - 770.","Schenkel E. (1902) Beitrag zur Kentnis der Dekapodenfauna von Celebes. Verhandlungen der Schweizerischen Naturforschenden Geselschaft (Basel, 13, 485 - 585. pls. 7 - 13.","Rathbun M. J. (1910) The Danish Expedition to Siam 1899 - 1900. v. Brachyura. Kongelige Danske Videnskabernes Selskab, Naturvidenskabelige Matematiske Afhandlinger, 5 (4), 301 - 367.","Roux J. (1917) Crustaces (Expedition de 1903). Nova Guinea, 5, 589 - 621.","Stebbing T. R. R. (1917) The Malacostraca of Natal. Annals of the Durban Museum, 2, 1 - 33.","Syomons C. T. (1920) Notes on certain shore crabs. Spolia Zeylanica, 306 - 313.","Balss H. (1922) Ostasiatische Decapoden. iv. Die Brachyuren (Canceridae). Archive Fur Naturgeschichte, A (11), 94 - 166.","Boyce D. R. (1924) The calling crabs of Durban Bay, Uca annulipes (Milne-Edwards). South African Journal of Natural History, 4, 250 - 252.","Maccagno T. (1928) Crostacei decapodi. Le specie del genre Uca Leach conservate nel Regio Museo Zoologico di Torino. Bolletino dei Musei di Zoologia e di Anatomia Comparata della R. Universita di Torino, 41 (11), 1 - 52.","Sakai T. (1936) Report on the Brachyura collected by Mr. F. Hiro at Palao (Pelew) Islands. Science Reports of the Tokyo Bunrika Daigaku, (B) 2, 155 - 177.","Miyake S. (1936) A list of known species of Brachyura from Ishigakishima. Annotationes Zoologicae Japonenses, 15 (4), 506 - 513.","Panikkar N. K. & Aiyar R. G. (1937) The brackish-water fauna of Madras. Proceeding of Indian Academic of Science, 6 (B 5), 284 - 337.","Suvatti C. (1938) A check-list of aquatic fauna in Siam (excluding fishes). Bureau of Fisheries (Bangkok), pp. 116.","Chace F. A. (1942) Decapod Crustacea. Scientific results of a fourth expedition to forested areas in Eastern Africa. iii. Bulletin of the Museum of Comparative Zoology, Harvard College, 91, 185 - 233.","Vatova, A. (1943) Decapoda della Somalia. Thalassia, 6 (2), 1 - 37.","Buitendijk A. M. (1947) Crustacea Anornura and Brachyura. Zoological notes frorn Port Dickson, iii. Zoologische Mededeelingen, 28, 280 - 284.","Barnard K. H. (1950) Descriptive Catalogue of South African decapod Crustacea (Crabs and Shrimps). Annals of the South African Museum, 38, 837 pp.","Tweedie M. W. F. (1950) Grapsoid crabs from Labuan and Sarawak. Sarawak Museum Journal, 5 (2), 356 - 367.","Sankarankutty C. (1961) On Decapoda Brachyura from the Andaman and Nicobar Islands. 1. Families Portunidae, Ocypodidae, Grapsidae and Mictyridae. Journal of the Marine Biological Association of India, India, 3 (1, 2), 101 - 119.","Forest J. & Guinot D. (1961) Crustaces decapodes brachyoures de Tahiti et des Tuamotu. In: Expedition Francaise sur les Recifs Coralliens de la Nouvelle-Caledonie. Volume preliminaire. Fondation Singer-Polignac, Paris, xi + 195 pp., pls. 1 - 18.","Claude M. (1964) Check list of the Crustacea Brachyura (Crabs) recorded from Mauritius. The Mauritius Institute Bulletin, 5 (1), pp. 1 - 60.","Pillay K. K. & Nair N. B. (1971 a) The annual reproductive cycles of Uca annulipes, Portunus pelagicus and Metapenaeus affinis (Decapoda. Crustacea) from the south-west coast of India. Marine Biology, Berlin, 11 (2), 152 166.","Rajendran A. (1972) Distribution of Uca annulipes (Latreille) in the intertidal mudflats of the Vellar estuary. Proceedings of the Indian National Science Academy, 38 (5 - 6), 378 - 379.","Naiyanetr P. (1980) Crustacean Fauna of Thailand (Decapoda and Stomatopoda). Department of Biology, Faculty of Science, Chulalongkorn University of Bangkok, Thailand, 73 pp.","Macia A., Quincardete, I. & Paula, J. (2001) A comparison of alternative methods for estimating population density of the fiddler crab Uca annulipes at Saco Mangrove, Inhaca Island (Mozambique). Hydrobiologia, 449, 213 - 219.","Ng P. K. L. & Davie P. J. F. (2002) A checklist of the Bbrchyuran crabs of Phuket and Western Thailand. Phuket Marine Biological Center Special Publication, 23 (2), 369 - 384.","Skov M. W., Vannini M., Shunula J. P., Hartnoll R. G. & Cannicci S. (2002) Quantifying the density of mangrove crabs. Ocypodidae and Grapsidae. Marine Biology (Berlin), 141 (4), 725 - 732.","Lee P. S. & Lim, S. S. L. (2003) Distributional patterns and species density of a fiddler crab community (Uca annulipes and U. vocans) at Telok Assam, Bako National Park, Sarawak. Sarawak Museum Journal, 58 (79), 183 198.","Lim S. S. L. (2004) A comparative study of some mouthpart adaptations of Uca annulipes (H. Milne Edwards, 1837) and U. vocans (Linnaeus, 1758) (Brachyura, Ocypodidae) in relation to their habitats. Crustaceana, 77 (10), 1245 - 1251.","Litulo C. (2004) Fecundity of the pantropical fiddler crab Uca annulipes (H. Milne Edwards, 1837) (Brachyura. Ocypodidae) at Costa do Sol mangrove, Maputo Bay, southern Mozambique. Western Indian Ocean Journal of Marine Science, 3 (1), 87 - 91.","Lim S. S. L., Lee P. S. & Diong C. H. (2005) Influence of biotope characteristics on the distribution of Uca annulipes (H. Milne Edwards, 1837) and U. vocans (Linnaeus, 1758) (Crustacea, Brachyura, Ocypodidae) on Pulau Hantu Besar, Singapore. Raffles Bulletin of Zoology, 53 (1), 111 - 114.","Ghory F. S. & Siddiqui F. A. (2006) A comparative study of the first zoeal stage of Uca urvillei and Uca annulipes (Crustacea. Brachyura. Ocypodidae) reared in the laboratory. Turkish Journal of Zoology, 30 (2), 161 166.","Naiyanetr P. (2007) Checklist of crustacean fauna in Thailand (Decapoda, Stomatopoda, Anostraca, Myodocopa and Isopoda). Open Biodiversity Series, 19, 197 pp.","Shih H. - T, Kamrani E., Davie P. J. F & Liu M. Y. (2009) Genetic evidence for the recognition of two fiddler crabs, Uca iranica and U. albimana (Crustacea: Brachyura: Ocypodidae), from the northwestern Indian Ocean, with notes on the U. lactea species-complex. Hydrobiologia, 635, 373 - 382.","Pesta O. (1911) Crustacea. Botanische und Zoologische Ergebnisse einer Wissenschaftlichen Forschunsreise nach den Samoa Inseln, Dem Neuguinea-Arcihpel und den Salomonsinseln, von Marz bis Dezember 1905. Kaiserlich- Koniglichen Hof und Staatsdruckerei, Wien, 30 pp, figs. 5, pl. 3.","Tesch J. J. (1918) The Decapoda Brachyura of the Siboga Expedition. II. Goneplacidae and Pinnotheridae. Siboga Expeditie Monographie, 39 c 1 (84), 149 - 295, pls. 7 - 18.","Bott R. (1973) Die Verwandtschaftliche Beziehungen de Uca - Arten (Decapoda: Ocypodidae). Senckenbergiana Biologica, 54, 315 - 325.","Dai Ai-Yun & Yang S. (1991) Crabs of the China Seas. China Ocean Press Beijing; Springer Berlin, Heidelberg, New York, Tokyo, 682 pp., pls. 1 - 74 (English edition).","Crane J. (1975) Fiddler Crabs of the World. Ocypodidae: Genus Uca. Princeton University Press, Princeton, New Jersey, i - xxiii + 736 pp.","Frith D. W. (1977) A preliminary list of macrofauna from a mangrove forest and adjacent biotops at Suran Island, western Peninsula Thailand. Phuket Marine Biological Center Research Bulletin, 17, pp. 1 16.","Vannini M. & Valmori P. (1981) Researches on the coast of Somalia. The shore and the dune of Sar Uanle. 31. Ocypodidae and Gecarcinidae. Monitore Zoologico Italiano, n. s., suppl. 14, 199 - 226.","Pretzmann G. (1971) Ergebnisse einiger Sammelreisen nach Vorderasien. 2. Teil: Marine Brachyura. Annalen des Naturhistorischen Museums Wien, 75, 477 - 487.","Kossmann R. (1877) Malacostraca (1. Teil: Brachyura). Zoologische Ergebnisse einer im Auftrage der Koniglichen Academie der Wissenschaften u Berlin ausgefuhrten Reise in die Kustengebiete des Roten Meeres, 3 (1), 66 pp, pls 1 - 3.","Rathbun M. G. (1924) Results of Dr. E. Mjoberg's Swedish scientific expeditions to Australia 1910 - 1913. 37. Brachyura, Albuneidae and Porcellanidae. Arkiv for Zoologi, 16 (23), 1 - 33."]}