Diversity and Distribution of Fiddler Crabs (Crustacea: Brachyura: Ocypodidae) around the Arabian Sea

© 2022 Academia Sinica, Taiwan Open Access Special issue: Systematics and Biogeography of Fiddler Crabs Diversity and Distribution of Fiddler Crabs (Crustacea: Brachyura: Ocypodidae) around the Arabian Sea Hsi-Te Shih1,* , Mani Prema2,§ , Austin Anand Jeya Kumar3,§ , Noor Us Saher4,§ , Samuthirapandian Ravichandran2, Sahir Odhano5, and Gustav Paulay6 1Department of Life Science and Research Center for Global Change Biology, National Chung Hsing University, Taichung 402, Taiwan. *Correspondence: E-mail: [email protected] (Shih) 2Centre of Advanced Study in Marine Biology, Annamalai University, Parangipettai - 608 502, Tamil Nadu, India. E-mail: [email protected] (Prema); [email protected] (Ravichandran) 3Department of Marine Biology, Faculty of Marine Science, King Abdulaziz University, Jeddah 21589, Saudi Arabia. E-mail: [email protected] (Kumar) 4Center of Excellence in Marine Biology, University of Karachi, Karachi 75270, Pakistan. E-mail: [email protected] (Saher) 5Department of Fisheries and Aquaculture, Shaheed Benazir Bhutto University of Veterinary and Animal Sciences, Sakrand67210, Sindh, Pakistan. E-mail: [email protected] (Odhano) 6Florida Museum of Natural History, University of Florida, Gainesville, FL 32611, USA. E-mail: [email protected] (Paulay) §MP, AJK, and NUS contributed equally to this paper. Received 1 January 2022 / Accepted 25 March 2022 / Published 16 November 2022 Communicated by Benny K.K. Chan Special issue: Systematics and Biogeography of Fiddler Crabs (articles 64–71). Editors: Hsi-Te Shih and Benny K. K. Chan Nine species of fiddler crabs (Crustacea: Ocypodidae: Gelasiminae) are known from the Arabian Sea and adjacent waters (Red Sea, Gulf of Aden, Gulf of Oman and Arabian/Persian Gulf): five species of Austruca, one Cranuca, two Gelasimus and one Tubuca. COI sequence data match morphological species boundaries and shows high connectivity within each. The fauna is highly endemic, with three species of Austruca (A. albimana, A. iranica and A. sindensis) confined to this region, and four others restricted to the Indian Ocean. restricted to the Indian Ocean. Austruca albimana and A. iranica speciated locally and now have narrowly overlapping ranges in Oman. Our results confirm the westernmost distributions of Austruca annulipes and Tubuca alcocki are Pakistan and the Red Sea, respectively. A key for the nine species is also provided to help identification. Key words: Fiddler crabs, Austruca, Cranuca, Gelasimus, Tubuca, Mitochondrial cytochrome c oxidase subunit I (COI), Barcodes. BACKGROUND Spanning more than half of the tropics, the IndoWest Pacific (IWP) is the largest and most diverse marine biogeographic region, recognized as a unit partly because many species range across its vast extent, from East Africa to the remote islands of Polynesia. Understanding diversification in this vast region relies in part on documenting the ranges, connectivity, and relationship of species. Genetic evidence is especially illuminating and reveals narrower ranges and cryptic diversity in some taxa, while confirming region-wide panmixia in others (Bickford et al. 2007; Teske et al. 2008; Lourenço et al. 2017). A major task today is to reevaluate species diversity and distributions established via morphology in the part, using integrative taxonomic methods. The Western Indian Ocean (WIO) is perhaps the most heterogeneous part of the IWP, and genetic scrutiny, especially, has led to the recognition of Citation: Shih HT, Prema M, Kumar AAJ, Saher NU, Ravichandran S, Odhano S, Paulay G. 2022. Diversity and distribution of fiddler crabs (Crustacea: Brachyura: Ocypodidae) around the Arabian Sea. Zool Stud 61:65. doi:10.6620/ZS.2022.61-65. Zoological Studies 61:65 (2022) doi:10.6620/ZS.2022.61-65 1 © 2022 Academia Sinica, Taiwan INDIA PAKISTAN IRAN IRAQ KUWAIT SAUDI ARABIA UAE OMAN YEMEN SOMALIA GulfofAden GulfofOman Arabian/PersianGulf Arabian Sea RedSea LaccadiveSea Indian Ocean SRI LANKA DJIBOUTI ERITREA EGYPT SUDAN ISRAEL JORDAN 1 5 4 3 2 678 9 10 11 12 15 14 13 ETHIOPIA greater diversity and more complex distributional patterns (DiBattista et al. 2015). The Arabian Sea and adjacent waters (Red Sea, Gulf of Aden, Gulf of Oman, Arabian/Persian Gulf) (Fig. 1) stand out as especially complex, with great regional and seasonal fluctuations in temperature, productivity, oxygenation, etc. This regional heterogeneity has led to substantial differences in the biota across this relatively small area, and both regional and local endemism, especially in the Red Sea, Gulf of Aden, and Oman (DiBattista et al. 2016; Wehe and Fiege 2002; Wells 2000). Currently 49 species of fiddler crabs are recognized in the IWP region (Crane 1975; Shih et al. 2016 2018 2019; Shih and Poupin 2020), and the diversity is as high as 23 species around the Coral Triangle (also known as the Malay Archipelago or the Indo-Australian Archipelago = IAA; Lohman et al. 2011). However, only 13 species are recorded from the WIO (from eastern Africa to western Indian subcontinent), with an additional questionable record of Tubuca paradussumieri (Bott, 1973) from Madagascar (Shih et al. 2018: 49). The genus Cranuca and seven species, viz. Austruca albimana (Kossmann, 1877), A. iranica (Pretzmann, 1971), A. occidentalis (Naderloo, Schubart & Shih, 2016), A. sindensis (Alcock, 1900), Cranuca inversa (Hoffmann, 1874), Paraleptuca chlorophthalmus (H. Milne-Edwards, 1837) and Tubuca urvillei (H. Milne-Edwards, 1852), are endemic to this area. In the northern part of the WIO, the Arabian Sea (including the coastal areas of Yemen, Oman, Pakistan and western India) and adjacent waters (including the Red Sea, Gulf of Aden, Gulf of Oman, and Arabian/ Persian Gulf) (Fig. 1), nine species of fiddler crabs have been reported, viz. Austruca albimana, A. annulipes (H. Milne-Edwards, 1837), A. iranica, A. sindensis, A. variegata (Heller, 1862), Cranuca inversa, Gelasimus hesperiae (Crane, 1975), G. tetragonon (Herbst, 1790) and Tubuca alcocki Shih, Chan & Ng, 2018 (Shih et al. 2009 2018 2021; Naderloo et al. 2010; Saher et al. Fig. 1. Collection sites for specimens in this study (filled circles) and an additional literature record (empty circle) (see Table 1). Dash line on the Indian Ocean represents the border of the Arabian Sea. page 2 of 18Zoological Studies 61:65 (2022) © 2022 Academia Sinica, Taiwan 2014; Naderloo 2017). Most studies of fiddler crabs in this region have focused on taxonomy and distribution, with a few on molecular phylogenetics (e.g., Shih et al. 2009 2018), population genetics (Shih et al. 2015; Odhano et al. 2018), larval development (Hashmi 1968; Ghory and Siddiqui 2006; Al-Aidaroos 2013), behavior and ecology (Mokhlesi et al. 2011; Saher and Qureshi 2012 2017). Studies of fiddler crabs around the Arabian Sea are comparatively few compared with other areas of the IWP, especially East Asia, Southeast Asia and Australia. In this study we assess each species in this fauna based on combined morphological and genetic evidence, update their distribution based on the literature and new specimens, and provide a key to the fauna. We also discuss the genetic connectivity and origin of the fauna. MATERIALS AND METHODS Specimens of fiddler crabs collected from the Arabian Sea and adjacent waters (including the Red Sea, Gulf of Aden, Gulf of Oman and Arabian/ Persian Gulf) (Table 1; Fig. 1) were preserved in 70% to 95% ethanol and deposited into the Centre of Advanced Study in Marine Biology, Annamalai University (CASAU), Parangipettai, Tamil Nadu, India; the Zoological Collections of the Department of Life Science, National Chung Hsing University, Taichung, Taiwan (NCHUZOOL) and the Zoological Reference Collection, Raffles Museum of Biodiversity Research, National University of Singapore (ZRC). Specimens deposited in Muséum National d'Histoire naturelle, Paris, France (MNHN); Museo Zoologico dell’Università di Firenze, Italy (MZUF); the Naturhistorisches Museum, Wien, Austria (NHMW); Steinhardt National Collections of Natural History, Tel Aviv University, Israel (TAU); and Florida Museum of Natural History, University of Florida, Florida, USA (UF) were also examined. The following abbreviations are used: CW = carapace width, CL = carapace length. In the synonymy of each species, only the references to the original description, relevant revisions and records from around the Arabian Sea are included. Genomic DNA was isolated from muscle tissue using kits (see Shih et al. 2016 for details). A portion of the COI gene was amplified with polymerase chain reaction (PCR) using the primers LCO1490, HCO2198 (Folmer et al. 1994), COL14 (Roman and Palumbi 2004), COL6, COH6 (Schubart 2009) and ICOUB (Huang et al. 2021), as well as the newly designed primers LCOB (5'-CAAAYCATAAAGAYATYGG-3'), HCOex (5'-GCTCATACTACAAATCCTAAA-3'), HCOex2 (5'-GCTCANACTACAAATCCTAA-3') and HCOex3 (5'-GCTCANACTACRAATCCTA-3'). PCR conditions for the above primers were 40 cycles of denaturation for 50 s at 94°C, annealing for 70 s at 45–47°C, and extension for 60 s at 72°C, followed by extension for 10 min at 72°C. Sequences were obtained by automated sequencing (Applied Biosystems 3730, USA) and deposited into GenBank (Table 1). Additional sequences of COI from related A. occidentalis from eastern Africa as well as A. variegata from the eastern Indian Ocean were downloaded from GenBank or sequenced from specimens with the same methods. A neighbor-joining (NJ) tree was generated with MEGA (vers. 11, Tamura et al. 2021), using the Kimura (1980) 2-parameter (K2P) model with the complete deletion option. K2P distances among specimens were also calculated in MEGA. RESULTS Molecular analyses of COI Specimens of eight of the nine species of fiddler crabs recorded from the region were represented among the specimens studied, and COI sequences were successfully generated from 65 specimens representing all eight species (Table 1, Fig. 4). Morphological and genetic identifications were fully congruent; there was no evidence for cryptic species in the region. Intra-specific variation was small, < 0.8% (K2P) for all species. There was no pattern of geographic differentiation across the areas sampled, including samples from the eastern Indian Ocean and western Pacific Ocean, for species for which samples were available from there. Interspecific divergences were all > 10%, lowest between G. hesperiae and G. tetragonon. TAXONOMY Family Ocypodidae Rafinesque, 1815 Subfamily Gelasiminae Miers, 1886 (sensu Shih et al. 2016) Genus Austruca Bott, 1973 Austruca albimana (Kossmann, 1877) (Fig. 2A, B) Gelasimus annulipes var. albimana Kossmann, 1877: 53 (type locality: Red Sea); Crane 1975: 298. Uca annulipes – Nobili 1906a: 312; Nobili 1906b: 150; Maccagno 1928: 35, fig. 20; Hornby 1997: 14 [part?]. Uca (Celuca) lactea annulipes – Crane 1975: 298, 611 [part]; Lewinsohn 1977: 61. page 3 of 18Zoological Studies 61:65 (2022) © 2022 Academia Sinica, Taiwan Uca lactea albimana – Price et al. 1987: 456. Uca albimana – Yamaguchi 1994: 153; Shih et al. 2009: 376; Shih et al. 2013: 643. Uca annulipes albimana – Apel and Türkay 1999: 133; Simões et al. 2001: 86. Uca (Paraleptuca) albimana – Ng et al. 2008: 241. Uca (Austruca) albimana – Naderloo et al. 2010: 4, figs. 1a–l, 4a, 6a, b; Naderloo et al. 2015: 409; El-Sayed et al. 2016: 7, pls. 3A, B, Table 1. Specimens and COI haplotypes of the fiddler crabs from the Arabian Sea and adjacent waters. The numbers within brackets following localities correspond to those in figure 1. *, sequences are shorter and not included for further analyses (see “Molecular analyses of COI”). See MATERIALS AND METHODS for abbreviations of museums and universities Locality Sample size Catalogue no. of NCHUZOOL (unless indicated) Austruca albimana Egypt: Sinai: Nabq - El Arwashie; Ras Mohammed [1] 2 13242 Saudi Arabia: Al Darb [4] 2 15002, 15003 Oman: Bar Al Hikman Peninsula [9] 1 UF7790 Austruca annulipes Pakistan: Sandspit: backwater mangrove area [11] 3 15088 India: Dongri, Uttan, Mira Bhayandar, Maharashtra [14] 1 15086 India: Mumbai [14] 1 15087 Thailand: Phuket 1 13257 Malaysia: Tioman 1 13243 China: Sanya, Hainan 1 13244 Austruca iranica Oman: Bar Al Hikman Peninsula [9] 2 UF7791, UF7752 Iran: Gavbandi, Hormozgan [6] 2 13245, 13247 Iran: Qeshm I., Hormozgan [8] 2 13246, 13248 Pakistan: Sandspit, Karachi, Sindh [11] 2 15091 Austruca occidentalis South Madagascar 1 ZRC THH04-30 South Madagascar 1 ZRC THH04-30 Kenya: Ngomeni Bay 1 MZUF 4297 Madagascar 1 UF14433 Austruca sindensis Iraq: Khur Al-Zubair [5] 2 ZRC 2010.0103 Iran: Bandar Abbas [7] 6 13645 Iran: Qeshm I., Hormozgan [8] 4 13576 Pakistan: Sonmiani, Balochistan [10] 1 13643; 13644 Pakistan: Sandspit, Karachi, Sindh [11] 4 13641 Pakistan: Sandspit, Karachi, Sindh [11] 6 13642 Austruca variegata India: Tamil Nadu 3 ZRC 2001.0853 India: Tamil Nadu 3 ZRC 2001.0853 India: Tamil Nadu 3 ZRC 2018.1375 India: West Bengal 2 ZRC 2017.0917 Cranuca inversa Egypt: Sinai: Nabq [1] 1 TAU SLR 1475 Egypt: Sinai: Nabq - El Arwashie; Ras Mohammed [1] 1 13623 Saudi Arabia: Al Darb [4] 2 15093 Somalia: Giumbo 1 MZUF 711 Kenya: Gazi 1 MZUF 1024 Tanzania: Dar es Salaam 1 13255 Gelasimus hesperiae Djibouti [3] 1 MNHN-IU-2011-5608 W India: Karnataka: Karwar: Kali river sangam [15] 4 15094, 15095 Thailand: Phuket 2 ZRC 2000.1056, ZRC 2001.1071 Gelasimus tetragonon Egypt: Sinai: Nabq - El Monqata [1] 1 15096 Eritrea: Dahlak Archipelago [2] 1 TAU EGg 3972 Madagascar: Belaza, Sarodrano 1 ZRC THH04-17 Taiwan: Penghu 1 13304, 14678 Tubuca alcocki Saudi Arabia: Al Darb [4] 2 14904 Pakistan: Sandspit [11] 3 15097 India: Diu [13] 1 14900 India: Mumbai [14] 6 14899, 14901, 14902, 14903, 14925 W India: Karwar: Sunkeri [15] 3 15098, 15099 Thailand: Ranong 1 ZRC 2017.1278 (holotype) Total 78 page 4 of 18Zoological Studies 61:65 (2022) © 2022 Academia Sinica, Taiwan 4A, D, G, J. Austruca albimana – Shih et al. 2016: 153, 168; Naderloo 2017: 412, figs. 37.10, 37.13a; Baakdah 2018: 51; AAJ Kumar 2019: 53, figs. 2g–i, 3a–d; Sasaki 2019: 12423. Austruca (Austruca) albimana – Rosenberg 2019: 734. Material examined: Egypt: 2 ♂♂ (15.3, 15.2 mm) (NCHUZOOL 13242), Nabq-El Arwashie, Sinai, Red Sea, coll. S. Barbaresi et al., 14 Oct. 2004. Saudi Arabia: 3 ♂♂ (10.4–12.1 mm) (NCHUZOOL 15085), 1 ♀ (9.7) (NCHUZOOL 15002), 1 ♀ (11.8 mm) (NCHUZOOL 15003), Al Darb (17°26'53.4"N, 42°17'03.3"E), coll. A. J. Kumar, 25 Apr. 2017. Oman: 1 ♂ (11.8 mm) (UF 7790), Bar al Hikman Peninsula, coll. V. Bonito and G. Paulay, 22–24 Jan. 2005; 1 ♂ Locality Haplotype Access. no. Austruca albimana Egypt: Sinai: Nabq - El Arwashie; Ras Mohammed [1] AaL1 AB471906 Saudi Arabia: Al Darb [4] AaL3, AaL4 ON075591, ON075592 Oman: Bar Al Hikman Peninsula [9] AaL2 ON075593 Austruca annulipes Pakistan: Sandspit: backwater mangrove area [11] Aan1, Aan2, Aan3 ON075594, ON075595, ON075596 India: Dongri, Uttan, Mira Bhayandar, Maharashtra [14] Aan4 ON075597 India: Mumbai [14] Aan5 ON075598 Thailand: Phuket Aan2 AB491160 Malaysia: Tioman Aan6 AB471907 China: Sanya, Hainan Aan6 AB471907 Austruca iranica Oman: Bar Al Hikman Peninsula [9] Ai1 AB471909 Iran: Gavbandi, Hormozgan [6] Ai2, Ai3 AB471908, AB471910 Iran: Qeshm I., Hormozgan [8] Ai1, Ai4 AB471909, AB471911 Pakistan: Sandspit, Karachi, Sindh [11] Ai5, Ai6 ON075599, ON075600 Austruca occidentalis South Madagascar Ao1 AB813669 South Madagascar Ao1 ON075601 Kenya: Ngomeni Bay Ao2 ON075602 Madagascar Ao2 ON075603 Austruca sindensis Iraq: Khur Al-Zubair [5] As2 LC015060 Iran: Bandar Abbas [7] As1, As2, As5 AB813673, LC015060, LC015063 Iran: Qeshm I., Hormozgan [8] As1, 2 AB813673, LC015060 Pakistan: Sonmiani, Balochistan [10] As1, As2, As3, AB813673, LC015060, LC015061 Pakistan: Sandspit, Karachi, Sindh [11] As1, As2, As3, As4, AB813673, LC015060, LC015061, LC015062 Pakistan: Sandspit, Karachi, Sindh [11] As1, , As2 AB813673, LC015060 Austruca variegata India: Tamil Nadu Av1 LC465131 India: Tamil Nadu Av2, Av3, Av4 LC465140, LC465141, LC465142 India: Tamil Nadu Av5 LC465143, LC465144, LC465145 India: West Bengal Av3, Av6 LC465146, LC465146 Cranuca inversa Egypt: Sinai: Nabq [1] Ci1 LC087973 Egypt: Sinai: Nabq - El Arwashie; Ras Mohammed [1] Ci2 ON07560 Saudi Arabia: Al Darb [4] Ci3, Ci4 ON075605, ON075606 Somalia: Giumbo Ci4 ON075607 Kenya: Gazi Ci4 AB813674 Tanzania: Dar es Salaam Ci5 AB471917 Gelasimus hesperiae Djibouti [3] —* ON075608 W India: Karnataka: Karwar: Kali river sangam [15] Gh1, Gh2, Gh3, Gh4 ON075609, ON075610, ON075611, ON075612 Thailand: Phuket Gh5, Gh2 AB535422, AB535423 Gelasimus tetragonon Egypt: Sinai: Nabq - El Monqata [1] Gt1 ON075613 Eritrea: Dahlak Archipelago [2] Gt2 ON075614 Madagascar: Belaza, Sarodrano Gt3 LC053377 Taiwan: Penghu Gt1 AB535431 Tubuca alcocki Saudi Arabia: Al Darb [4] Ta1, Ta2 ON075615, ON075616 Pakistan: Sandspit [11] Ta1 ON075617, ON075618, ON075619 India: Diu [13] —* ON075620 India: Mumbai [14] Ta1 LC150445 W India: Karwar: Sunkeri [15] Ta1 ON075621, ON075622, ON075623 Thailand: Ranong Ta1 LC150445 Total Table 1. (Continued) page 5 of 18Zoological Studies 61:65 (2022) © 2022 Academia Sinica, Taiwan (17.2 mm) (UF 7789), Bar Al Hikman Peninsula, coll. V. Bonito and G. Paulay, 22–24 Jan. 2005; 1 ♂ (not measured) (UF 58686), Bandar Al Khiran sand flat (23.50856°N, 58.73186°E), coll. S. Brown, D. Uyeno, G. Koblasova and M. Cherneva, 26. Jan. 2020. Distribution: Egypt (Red Sea), Saudi Arabia (Red Sea), Eritrea, Djibouti, Yemen (including Socotra), Oman and UAE. Remarks: Because the Red Sea is semi-enclosed, Austruca albimana was thought to be endemic to this area (Shih et al. 2009). However, Naderloo et al. (2010) and Naderloo (2017) extended its distribution to the southern coast of the Gulf of Oman and the Arabian/ Persian Gulf (Oman and UAE). Comparisons of COI sequences from specimens from central Oman (Barr al Hikman) and the Red Sea demonstrate genetic connectivity across this area (Fig. 4). Austruca annulipes (H. Milne Edwards, 1837) (Fig. 2C, D) Cancer vocans minor Herbst, 1782: 81, pl. 1 (10) [nomen oblitum]; Shih et al. 2021: 208, fig. 1A. Gelasimus annulipes H. Milne Edwards, 1837: 55, pl. 18 (10–13) (type locality: mer des Indes” (= Indian Ocean)) [nomen protectum]; Kossmann 1877: 53; Kingsley 1880: 148, pl. 10(22) [part]; Alcock 1900: 353; Chhapgar 1957: 508, pl. 13j–o. Uca (Celuca) lactea annulipes – Crane 1975: 298, 611, figs. 18A–C, 19I–N, 20D–F, 24N–O, 27I, J, 29D, 32L, M, 41A, 54I, II, 69D, pls. 39A–D, 40C, D [part]. Uca annulipes – Tirmizi and Ghani 1996: 105, fig. 40; Shih et al. 2009: 376; Shih et al. 2013: 643; Saher et al. 2014: 672. Uca (Paraleptuca) annulipes – Ng et al. 2008: 241. Uca (Austruca) annulipes – Naderloo et al. 2010: 7, figs. 2a–h, 3a–e, 4b, 12a–c; Trivedi et al. 2015: 27. Uca lactea annulipes – Trivedi et al. 2012: 17, 20, fig. 3c; Beleem et al. 2014: 420, 422, pl. 2. Austruca annulipes – Shih et al. 2016: 153, 168, fig. 8A; Trivedi et al. 2018: 53; Beleem et al. 2019: 15; Sasaki 2019: 12424; Shih et al. 2021: 212, fig. 2; Pati et al. 2022: 526, fig. 12A–D. Austruca lactea – Beleem et al. 2019: 15. Austruca (Austruca) annulipes – Rosenberg 2019: 734. Material examined: Pakistan: 3 ♂ ♂ (15.5–20.8 mm) (NCHUZOOL 15088), backwater mangrove area, Sandspit, Karachi, coll. N. U. Saher, 10 Dec. 2020. India: 2 ♂♂ (13.5, 14.6 mm) (ZRC 2016.0191), Narayan Sarovar, Gulf of Kachchh, Gujarat, coll. 26 Mar. 2015; 12 ♂♂ (7.2–14.7 mm), 8 ♀♀ (5.6–9.8 mm), 1 ovig. ♀ (11.4 mm) (NCHUZOOL 15086), Dongri, Uttan, Mira Bhayandar, Maharashtra, coll. H.-N. Chen et al., 16 Mar. 2010; 6 ♂♂ (8.2–10.6 mm), 2 ♀♀ (6.5, 9.6 mm) (NCHUZOOL 15087), Mumbai, coll. H.-N. Chen et al., 17 Mar. 2010. Distribution: Pakistan, India, Myanmar, Thailand, Malay Peninsula, Singapore, Indonesia, Borneo, Vietnam, and China (Hainan). Remarks: Austruca annulipes was thought to be widely distributed in the IWP (Crane 1975; Naderloo et al. 2010). However, Shih et al. (2013) showed the specimens from eastern Africa represent a different species (as “Uca aff. annulipes”), later described as Uca occidentalis Naderloo, Schubart & Shih, 2016 (Naderloo et al. 2016). COI sequences show minimal genetic divergence suggesting good connectivity across the remainder of the species’ range from Pakistan to China (Fig. 4). Saher et al. (2014) noted that A. iranica and A. annulipes are sympatric in Pakistan, confirmed with COI data here. Austruca iranica (Pretzmann, 1971) (Fig. 2E, F) Gelasimus lacteus – Alcock, 1900: 355. Uca annulipes – Stephensen 1946: 189; Pretzmann 1971: 481; AlGhais and Cooper 1996: 419, figs. 7, 8; Hornby 1997: 14 [part?]; Tirmizi and Ghani 1996: 105 [part; not fig. 40]; Mokhlesi et al. 2011: 245. Uca annulipes iranica Pretzmann 1971: 481, pl. 5, figs. 11, 12 (type locality: Bandar-Abbas, Iran). Uca (Celuca) lactea annulipes – Crane 1975: 298, 611 [part]. Uca annulipes iranica – Apel and Türkay 1999: 133. Uca iranica – Shih et al. 2009: 376; Naderloo et al. 2013: 450; Shih et al. 2013: 643. Uca (Austruca) iranica – Naderloo et al. 2010: 14, figs. 6c–f, 7a, 8a, 9a–k; Naderloo and Türkay 2012: 54; Saher et al. 2014: 669, figs. 1–4; Naderloo et al. 2015: 409. Austruca iranica – Shih et al. 2016: 153, 168, fig 8C; Naderloo 2017: 413, figs. 37.11, 37.13b; Trivedi and Vachhrajani 2017: 82, figs. 4, 5; Trivedi et al. 2018: 54; Beleem et al. 2019: 15; Sasaki 2019: 12429; Pati et al. 2022: 527. Austruca (Austruca) iranica – Rosenberg 2019: 734. Material examined: Oman: 1 ♂ (13.8 mm) (UF 7791), Bar Al Hikman Peninsula, coll. V. Bonito and G. Paulay, 22–24 Jan. 2005; 1 ♀ (10.8 mm) (UF 7752), Bar Al Hikman Peninsula, coll. V. Bonito et al., 24 Jan. 2005. Iran: 1 ♂ (16.4 mm) (NCHUZOOL 15090), Qeshm I., Hormozgan Prov., coll. Jun. 2008; 2 ♂♂ (12.4, 14.0 mm), 1 ♀ (13.8 mm) (NCHUZOOL 13245), 1 ovig. ♀ (12.1 mm) (NCHUZOOL 13247), Gavbandi, Hormozgan, coll. E. Kamrani, Jul. 2008; 3 ♂♂ (13.4–16.4 mm) (NCHUZOOL 13248), 1 ♀ (11.0 mm) (NCHUZOOL 13246), Qeshm I., Hormozgan Prov., coll. E. Kamrani, Jul. 2008; 6 ♂♂ (12.8–15.7 mm), 2 ♀♀ (12.4, 13.9 mm) (NCHUZOOL 15089), Dokohak, Qeshm I, Hormozgan Prov., coll. E. Kamrani, 23 Jan. 2013. Pakistan: 1 ♂ (18.7 mm), 1 ♀ (15.2 mm) (NCHUZOOL 15091), Sandspit, Karachi, Sindh, coll. 6 Mar. 2013. Distribution: Oman, Iran, Pakistan and northwestern India. Remarks: Although A. iranica was thought to be restricted to the Arabian/Persian Gulf (Shih et al. 2009), page 6 of 18Zoological Studies 61:65 (2022) © 2022 Academia Sinica, Taiwan Fig. 2. Dorsal view of carapace and frontal view of major cheliped. Austruca albimana (A, B, NCHUZOOL 13242, CW 15.2 mm, Egypt); A. annulipes (C, NCHUZOOL 13642, CW 17.4 mm, Pakistan; D, NCHUZOOL 15088, CW 19.4 mm, Pakistan); A. iranica (E, F, NCHUZOOL 15090, CW 16.4 mm, Iran); A. sindensis (G, H, NCHUZOOL 13642, CW 17.4 mm, Pakistan). page 7 of 18Zoological Studies 61:65 (2022) © 2022 Academia Sinica, Taiwan this is unlikely as the Gulf is young and was dry land during the last glacial maximum at 18,000 year BP (Lambeck 1996). Later studies showed its distribution extended eastward to the Gulf of Oman, Pakistan and northwestern India (Naderloo et al. 2010; Saher et al. 2014; Naderloo 2017; Trivedi and Vachhrajani 2017). Here we record this species from the Arabian Sea coast of Oman, a new western distributional limit, where it co-occurs in the same coastal mangrove with A. albimana. COI sequence data shows good connectivity across the distribution of the species (Fig. 4). Austruca sindensis (Alcock, 1900) (Fig. 2G, H) Gelasimus inversus var. sindensis Alcock 1900: 356 (type locality: Karachi, Pakistan). Uca (Amphiuca) inversa sindensis – Crane 1975: 107, 599, figs. 69j, pl. 16E–H. Uca sindensis – Collins et al. 1984: 318, figs. 2, 3a–e, h, 4a, b, f, h; Apel and Türkay 1999: 134; Mokhlesi et al. 2011: 245; Saher and Qureshi 2012: 124; Naderloo et al. 2013: 450; Shih et al. 2013: 643; Beleem et al. 2014: 422; Saher et al. 2014: 672. Uca (Paraleptuca) sindensis – Ng et al. 2008: 241; Naser et al. 2010: e87 (pp. 1, 2), fig 2; Naderloo and Türkay 2012: 54; Naderloo et al. 2015: 409; Trivedi et al. 2015: 27. Austruca sindensis – Shih et al. 2016: 153, 168, fig 8G; Naderloo 2017: 414, figs. 37.12, 37.13c; Odhano et al. 2018: 313; Trivedi et al. 2018: 54; Beleem et al. 2019: 15; Sasaki 2019: 12439; Pati et al. 2022: 528. Austruca (Sinduca) sindensis – Rosenberg 2019: 734. Material examined: Iraq: 2 ♂♂ (16.1 mm, broken) (ZRC 2010.0103), Khur Al-Zubair. Iran: 1 ♀ (12.9 mm) (NCHUZOOL 15092), Qeshm I. Hormozgan, coll. Jun. 2008; 1 ♂ (16.3 mm), 3 ♀♀ (10.8–11.7 mm) (NCHUZOOL 13576), Qeshm I. Hormozgan, coll. E. Kamrani, Jul. 2008; 11 ♂♂ (6.9–12.3 mm) (NCHUZOOL 13645), Bandar Abbas, Hormozgan, coll. E. Kamrani, 3 Feb. 2013. Pakistan: 2 ♂♂ (12.1–12.9 mm), 2 ♀♀ (9.3–11.2 mm) (NCHUZOOL 13643), Sonmiani, Balochistan, coll. N. U. Saher, 14 May 2012; 8 ♂♂ (9.2–12.0 mm) (NCHUZOOL 13644), Sonmiani, Balochistan, coll. N. U. Saher, 9 Feb. 2013; 2 ♂♂ (18.7–19.0 mm), 2 ♀♀ (14.3–15.8 mm) (NCHUZOOL 13641), Sandspit, Karachi, Sindh, coll. N. U. Saher, 14 May 2012; 9 ♂♂ (10.2–17.4 mm) (NCHUZOOL 13642), Sandspit, Karachi, Sindh, coll. N. U. Saher, 6 Mar. 2013. Distribution: Kuwait, Iraq, Iran, Pakistan and western India. Remarks: Austruca sindensis was described from Karachi, Pakistan (Alcock 1900) and later reported from adjacent areas, including Kuwait, Iraq, Iran and western India (Shih et al. 2015; Trivedi et al. 2018; Beleem et al. 2019). There are no records of this species from the southern coasts of the Arabian/Persian Gulf and the Gulf of Oman (Naderloo 2017). Austruca variegata (Heller, 1862) Gelasimus variegatus Heller, 1862: 521 (type locality: Madras, India); Kingsley 1880: 154; Crane 1975: 326. Gelasimus perplexus – Heller 1865: 38, pl. 5(4); A. Milne-Edwards 1873: 274. Uca triangularis – Kappalli et al. 2012: 967; Supriya et al. 2017: 647. Uca (Celuca) triangularis bengali Crane 1975: 286, figs. 24I, J, 32N, O, 59A, 68C, 101 [part]. Austruca bengali – Shih et al. 2016: 153 [part]; Trivedi et al. 2018: 54. Austruca triangularis – Trivedi et al. 2018: 54 [part]. Cranuca inversa – Trivedi et al. 2018: 54. Austruca variegata – Sasaki 2019: 12442; Shih et al. 2019: 4, figs. 1a–c, 2, 3a, b, 4a–d, 5a–d, 6a–c, f, 7a, b, e, 8a–d, 9. Material examined: Lectotype, ♂ (CW 16.7 mm, PL 24.7 mm) (NHMW 25656; original Novara label, #74 = AN.1866.II.74.; labeled as “Gelasimus perplexus M. Edw.”), Madras, India, coll. Johann Zelebor. Distribution: Southwestern and eastern India, Sri Lanka and Bangladesh. Remarks: Austruca variegata ranges mostly along the Bay of Bengal. This species has been frequently misidentified, as A. bengali, A. perplexa, A. triangularis and Cranuca inversa (see Shih et al. 2019). Shih et al. (2019) resurrected this species with morphological and molecular evidence. Although no specimens from western Indian were examined for our study, this species is recorded from the area by Kappalli et al. (2012) and Supriya et al. (2017) as “Uca triangularis”, and its distribution appears to extend to the Laccadive Sea (Shih et al. 2019). Genus Cranuca Beinlich & von Hagen, 2006 Cranuca inversa (Hoffmann, 1874) (Fig. 3A, B) Gelasimus inversus Hoffmann 1874: 19, pl. 4 (23–26) (type locality: Nossi-Faly, Madagascar); Kingsley 1880: 155. Uca inversa – Pocock et al. 1903: 213, 216; Nobili 1906a: 312; Nobili 1906b: 151; Maccagno 1928: 26, fig. 13; Collins et al. 1984: 318, 324, figs. 3f, g, 4c–e, g; Yamaguchi 1994: 164; Al-Ghais and Cooper 1996: 421; Hornby 1997: 14; Apel and Türkay 1999: 134; Simões et al. 2001: 86; Hogarth and Beech 2002: 24, 1 unnumbered fig.; Naderloo et al. 2013: 450; Shih et al. 2013: 643. Uca (Amphiuca) inversa inversa – Crane 1975: 105, 599, figs. 39G, 69I, pl. 16A–D; Lewinsohn 1977: 63. Uca inversa inversa – Fransen et al. 1997: 150; Price et al. 1987: 456. Uca (Cranuca) inversa – Beinlich and von Hagen 2006: 10, fig. 5a; Ng et al. 2008: 240; Naderloo and Türkay 2012: 53; Naderloo et al. 2015: 409; El-Sayed et al. 2016: 9, pls. 3C, D, 4B, E, H, K. Cranuca inversa – Shih et al. 2016: 153, 169, fig. 9A; Naderloo 2017: 416, fig. 37.13d, 37.14; Baakdah 2018: 51; AAJ Kumar 2019: 54, figs. 2j, 3e–g; Rosenberg 2019: 734; Sasaki 2019: 12443. page 8 of 18Zoological Studies 61:65 (2022) © 2022 Academia Sinica, Taiwan Material examined: Egypt: 1 ♂ (17.4 mm) (TAU SLR 1475), Nabq, Sinai, coll. 19 May 1968; 1 ♀ (18.1 mm) (NCHUZOOL 13623), Nabq-El Arwashie, Sinai, coll. S. Barbaresi et al., 14 Oct. 2004. Saudi Arabia: 4 ♂♂ (16.5–19.9 mm) (NCHUZOOL 15093), Al Darb (17°26'53.4"N, 17°26'53.4"N), coll. A. J. Kumar, 25 Apr. 2017. Distribution: South Africa to Somalia, Madagascar, Egypt (Red Sea), Saudi Arabia (Red Sea), Eritrea, Yemen, Oman, UAE and Iran. Remarks: The southernmost record of this species from Natal, South Africa (Crane 1975) needs to be substantiated; confirmed records extend southward to Kosi Bay (near the Mozambican border) (Peer et al. 2015). The northernmost record is from Qeshm Island, Iran along the northern coast of the Arabian/Persian Gulf (Naderloo 2017). COI data confirm connectivity of populations between the Red Sea and the East African coast (Fig. 4). Genus Gelasimus Latreille, 1817 Gelasimus hesperiae (Crane, 1975) (Fig. 3C, D) ?Gelasimus tetragonon var. spinicarpa Kossmann, 1877: 52 (type locality: Red Sea); Crane 1975: 80; Rosenberg 2013: 494. Gelasimus cultrimanus – Kingsley 1880: 140, pl. 9(7) [part]. Gelasimus marionis – Kingsley 1880: 141, pl. 9(8) [part]. Uca Marionis – Nobili 1906a: 314. Uca Marionis var. nitida – Nobili 1906a: 314. Gelasimus marionis – Chhapgar 1957: 509, pl. 13p, q, s, t, v, w. Gelasimus marionis nitidus – Chhapgar 1957: 510, pl. 13r, u. Uca (Thalassuca) vocans hesperiae Crane, 1975: 89, 92, figs. 64E, EE (type locality: Zanzibar, Tanzania). Uca vocans – Yamaguchi 1994: 180. Uca vocans forma excisa – Al-Ghais and Cooper 1996: 421. Uca vocans hesperiae – Fransen et al. 1997: 152. Uca hesperiae – Apel and Türkay 1999: 134. Uca (Gelasimus) hesperiae – Ng et al. 2008: 240; Naderloo et al. 2015: 409. Uca (Gelasimus) vocans – Trivedi et al. 2015: 27. Gelasimus hesperiae – Shih et al. 2016: 151, 169, fig. 7C; Naderloo 2017: 416, figs. 37.13f, 37.16; Trivedi et al. 2018: 54; Sasaki 2019: 12447. Gelasimus vocans – Trivedi et al. 2018: 54; Beleem et al. 2019: 15; Pati et al. 2022: 528, fig. 13A, B. Gelasimus (Gelasimus) hesperiae – Rosenberg 2019: 734. Material examined: Eritrea: 2 ♂♂ (21.0, 22.0 mm), 1 major chela (SMF 5645), Sahil, Massaua, selten überspülte Schlickzone mit Salicornia-artigem Bewuchs (15°36'37.1"N, 39°28'16.1"E), coll. E. Rüppell, Jan.-Mar. 1827. Djibouti: 1 ♂ (18.5 mm) (MNHN-IU-2011-5608), 1 ♂ (20.3 mm) (MNHNIU-2011-5607), Mission Gravier, coll. 3 Feb. 1904. India: 4 ♂♂ (13.7–18.2 mm), 1 ovig. ♀ (15.8 mm) (NCHUZOOL 15094), 5 ♂♂ (14.2–19.4 mm), 3 ovig. ♀♀ (13.5–15.9 mm) (NCHUZOOL 15095), 3 ♂♂ (12.5–13.1 mm), 2 ♀♀ (9.3–8.5 mm) (CASAUCR-1012), Kali R. estuary, Karwar, Karnataka, coll. M. Prema, 4 Feb. 2021. Distribution: South Africa to Somalia, Eritrea, Djibouti, Yemen, Oman, UAE, India, Sri Lanka, Bangladesh, Myanmar, western Thailand, Malaysia (western Malay Peninsula), Singapore and Indonesia (West Sumatra). Remarks: Gelasimus tetragonon var. spinicarpa was established by Kossmann (1877: 52) on three males from the Red Sea. One syntype (Leiden Museum cat. no. 1493, CL 12 mm, major propodus 23 mm) was identified as a species of the Gelasimus vocans complex by Crane (1975: 80), although the other two syntypes were not examined (Fransen et al. 1997: 152). Based on location, these Red Sea specimens would fall within Crane’s G. vocans herperiae. Rosenberg (2013) called attention to a potential homonymy between Uca spinicarpa Rathbun, 1900 and Gelasimus tetragonon var. spinicarpa Kossmann, 1877, if both were in Uca. However, as these species are now considered to be in their original genera, no homonymy exists and G. spinimana is an available name, and potential senior synonym of G. hesperiae. Morphological examination and sequencing of Red Sea topotypical samples is needed to assess this synonymy. The type locality of Gelasimus hesperiae is Tanzania (Zanzibar), with additional records from Madagascar, Eritrea, Sri Lanka, western Malay Peninsula and even Sulawesi in Crane (1975: 597). In the WIO, its distribution is extended southward to South Africa (Mngazana) (Peer et al. 2015) and eastward to Mumbai, western India (Chhapgar 1957). COI data shows high connectivity between sampled areas in W India and Thailand. Gelasimus tetragonon (Herbst, 1790) (Fig. 3E, F) Cancer tetragonon Herbst 1790: 257, pl. 20 (110) (type locality: unknown). Gelasimus tetragonon – Kossmann 1877: 52; Kingsley 1880: 143, pl. 9(11); Shih et al. 2016: 151, 169, fig. 7F; Naderloo 2017: 418, figs. 37.13e, 37.18; Trivedi et al. 2018: 54; Sasaki 2019: 12450. Uca tetragonon – Nobili 1906a: 313; Nobili 1906b: 151; Maccagno 1928: 22, figs. 7, 8; Price et al. 1987: 456; Apel and Türkay 1999: 134. Uca (Thalassuca) tetragonon – Crane 1975: 77–81, 596, figs. 37D, 63A, B, 81F, 82E, pl. 13; Lewinsohn 1977: 59; Sakai 1999: 38, plate 20D (holotype). Uca tetragonon – Yamaguchi 1994: 177, 184. Uca (Gelasimus) tetragonon – Ng et al. 2008: 240; Naderloo et al. 2015: 409; El-Sayed et al. 2016: 10, pls. 3E, F, 4C, F, I, L. Gelasimus (Mesuca) tetragonon – Rosenberg 2019: 734. Material examined: Egypt: 2 ♂ ♂ page 9 of 18Zoological Studies 61:65 (2022) © 2022 Academia Sinica, Taiwan Ghory FS, Siddiqui FA. 2006. 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