The Brachyuran Crabs (Crustacea: Decapoda) of Hong Kong: a Historical Review and Catalogue

© 2024 Academia Sinica, Taiwan Open Access The Brachyuran Crabs (Crustacea: Decapoda) of Hong Kong: a Historical Review and Catalogue Kingsley J. H. Wong1,2 , Peter K. L. Ng3, Hsi-Te Shih4, and Benny K. K. Chan2,* 1Institute of Ecology and Evolutionary Biology, National Taiwan University, Taipei 106, Taiwan. E-mail: [email protected] (Wong) 2Biodiversity Research Center, Academia Sinica, Taipei 115, Taiwan. *Correspondence: E-mail: [email protected] (Chan) 3Lee Kong Chian Natural History Museum, Faculty of Science, National University of Singapore, Singapore 117377, Republic of Singapore. E-mail: [email protected] (Ng) 4Department of Life Science and Global Change Biology Research Center, National Chung Hsing University, Taichung 402, Taiwan. E-mail: [email protected] (Shih) Received 18 April 2023 / Accepted 19 July 2024 / Published 31 December 2024 Communicated by Ryuji Machida The brachyuran crab fauna of subtropical Hong Kong is documented and an updated catalogue presented of all known species. Starting with the milestone studies by William Stimpson in the 1850s, many generations of workers have since added to the knowledge of this fauna. In the previous summary of the fauna by Chia-Jui Shen in 1940, 187 species were reported. Eight decades later, we now report 382 species from 27 superfamilies and 49 families, of which 22 species are new records. The present paper also reviews the history of carcinology in Hong Kong. The taxonomy of each species is treated, and a complete scientific bibliography is presented as far as possible. Key words: Hong Kong, Brachyuran fauna, Historical review, Literature, Taxonomy, Biodiversity, New records BACKGROUND Hong Kong is a small city in southern China with a land area of only 1107 km2 and with coastal territorial waters of 1648 km2 (Lands Department, HKSAR 2020). The territory seas of Hong Kong, in general, are shallow, with depths less than 50 m on average. This subtropical environment, nevertheless, exhibits remarkable spatial and temporal biodiversity (Watts 1973; Morton and Morton 1983) and contains a wealth of marine organisms with species counts reaching 6,000 (TPT Ng et al. 2016). Brachyuran crabs of diverse forms are found in nearly all aquatic habitats, and total number of species currently exceeds 7,200 globally (Davie et al. 2015a). Elsewhere in East Asia, some 1,073 species were recorded from Chinese seas, 800 from Taiwan (Yang et al. 2008; Ng et al. 2017a), while over 1,250 from Japan (T Sakai 1976; K Sakai 2004; Sasaki 2020). S.Y. Lee and Leung’s (1999) estimate that the brachyuran species diversity of Hong Kong was in excess of 300 has proven to be correct. Herein we report 382 species, representing a third of the total Chinese fauna. The fauna of Hong Kong is predominately intertidal and shallow-water, and is represented by 27 superfamilies and 49 families; the most species-rich groups being the Grapsoidea (51 spp.), Ocypodoidea (49 spp.), Portunoidea (49 spp.) and Pilumnoidea (43 spp.). In addition to the rich habitats present in Hong Kong, this small locality has also benefited from a long history of exploration of its seas and natural history investigations. A key prelude was the North Pacific Exploring Expedition (1853–1856) of the United States in East Asia, which was followed by several workers who collected extensive materials that were eventually deposited in many major European and American Institutes. By 1940, a comprehensive checklist of 187 species was compiled by the Chinese taxonomist ChiaJui Shen (Shen 1940a). A detailed historical review of the local brachyuran fauna is provided in the present Citation: Wong KJH, Ng PKL, Shih HT, Chan BKK. 2024. The brachyuran crabs (Crustacea: Decapoda) of Hong Kong: a historical review and catalogue. Zool Stud 63:38. doi:10.6620/ZS.2024.63-38. Zoological Studies 63:38 (2024) doi:10.6620/ZS.2024.63-38 1 © 2024 Academia Sinica, Taiwan work below. For eight decades since 1940, the contributions by many workers have substantially increased the number of brachyuran taxa known from Hong Kong. A comprehensive compilation, however, has never been done until now. To facilitate future research in biodiversity and the associated fields of conservation and management, we have included detailed taxonomic and ecological information for each species whenever possible. HISTORICAL ACCOUNT AND GENERAL DISCUSSION Our understanding of the locality called “Hong Kong” has evolved over time since 1842. Chinese characters representing Hong Kong appeared in documented publications not later than the midlate Ming Dynasty in Yueh-Ta-Chi (printed in 1595), referring to a small remote island off the coast of present day Guangdong (for pre-colonial circumstances, see e.g., Hayes 1984), while the later British colony was established as a direct result of the Treaty of Nanking signed between the British and Qing China after the First Opium War (1839–1842). The north-south oriented Pearl River leads to Canton (now Guangzhou), a longtime nexus of inter-continental maritime trade, and the colonized island at the east of the estuarine seas was first referred as “Hong Kong” in international literature. The British territory later extended north to include the peninsula of Kowloon opposite the Harbour in 1860 under the Convention of Beijing, and later leased the area south of the Shenzhen (= Sham Chun) River for another 99 years in 1898 under the Convention for the Extension of Hong Kong Territory. Between these periods, the boundaries of “Hong Kong” expanded, from a single island (1840s–1859), to both sides of the Harbour (1860–1898), and eventually to include the land south of the Shenzhen River (Figs. 1, 2; in Fig. 2 territory of these three stages shown in different tones of gray). The boundaries of “Hong Kong” have confused some workers. For example, Shen (1935) described a new species, Camptandrium anomalum (now Shenius, Dotillidae), based on material from “Taipo, near Canton, China”. The site from which this mangrove-obligate species (Ng et al. 2010b) was collected was in fact in the present day northeastern New Territories along the Tolo Channel, Hong Kong. As such, records from localities previously listed as “Canton”, or “South China”, are actually within the present recognized territory of Hong Kong, are included in the present paper. In the present section, as part of our historical review, we elaborate on how knowledge of the local brachyuran fauna accumulated through the years, emphasis being given to works which contribute new species and records. The list of names of authors of Asian ethnic origins, as well as titles of older Chinese text mentioned, are listed under appendix 1. Fig. 1. Map showing main locations in Hong Kong: A, location of Pearl River Delta in East Asia; B, location of Hong Kong in eastern flank of Pearl River Delta; C, map of Hong Kong (territorial seas shaded in medium gray). page 2 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Fig. 2. Map of Hong Kong before most large-scale reclamation works. Parts of the colony, namely Hong Kong Island (1842–1997), Kowloon Peninsula (1860–1997) and the New Territories (1898–1997) are shaded in different tones of gray. Local sites surveyed by W. Stimpson in the 1850s and C.-J. Shen in the 1930s are shown respectively as circles and squares, those of which have undergone substantial changes and/or urbanization are shown by a crossed icon. page 3 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan A welcome station for circumnavigations (1850s to the early 20th Century) In older Chinese texts, various intertidal crabs had been recorded (see Dai and Yang 1991), while those mentioning sites in the vicinity of Guangdong, South China, emerged later in the mid-Ming Dynasty. These works include as Guangdong Tongzhi (or Gazetteer of the Guangdong Province: first edition in 1535), and Guangdong Xinyu (or New Essays of Guangdong: printed in 1687). Regrettably, these records cannot be used as reliable or formal records of the fauna of the region, as they largely treated crabs as local produce, occasionally providing remarks on their edibility and/or flavor. On the other hand, western (European and American) naturalists have long travelled to Asia for exploration along established trade routes, collecting specimens, reporting and describing new taxa. Specimens were either collected by the naturalists themselves, and/or passed to them by locals or colleagues. Under the British administration from the 1840s, the colony was open for foreign trade and residency, and in the decades that followed, the population grew from 7,450 in 1841, to nearly 87,000 by 1859 (SC Fan 1974). The prosperity attracted European settlers, among whom were many keen naturalists (F-t Fan 2004). The study of local terrestrial plants, for example, became the pioneer volume of Flora Hongkongensis (Bentham 1861). Understanding of invertebrates in the region, especially for brachyurans, however, only started from the circumglobal expeditions of the west. Most early knowledge of brachyurans can be traced to the works of American zoologist William Stimpson (1832–1872), who served as the on-board zoologist, or Chief Scientific Officer, of the North Pacific Exploring Expedition (NPEE, 1853–1856) of the United States. The NPEE was the second circumnavigation organized by the US, following the first, United States Exploring Expedition (USEE; 1838– 1842). Main intentions included securing trade routes, exploring new whaling grounds, and providing a strong platform for naturalists and scientists (e.g., see Wilkes 1845; Bryan 1940). Extensive details of the NPEE had been revealed in Stimpson’s journal, edited and published by Vasile et al. (2005). The Expedition spent nearly nine months in the vicinity of Hong Kong (19th March–11th September 1854, and 29th January–6th April 1855, interrupted by a brief northward visit to the Ryukyus and Japanese seas), collecting extensively by dredging, shore excursions and market visits. In Chinese waters, their team accessed numerous localities, such as outlying islands eastward of Lei-Yue Mun (narrow passage east of Victoria Harbour, then as Li-ee-moon), Tung Lung Chau (as Tam-too), Nine Pins, Waglan, Po Toi (as Pootoy) and Dangan Dao (as Grand Lema), and westwards at Kap Shui Mun (as Cum-sing-moon) and Macao (see maps reproduced in Figs. 2 and 3). In a series of preliminary papers, Stimpson (1857 1858a b c d) recorded 287 brachyuran species, of which 126 were new to science. More details and illustrations were provided in his posthumously-published compendium of 1907. Of these taxa, 110 (including three variants of Chlorodius exaratus, Xanthidae, cf. Stimpson 1907) were reported from seas of South China, most (97 species and two variants) of which were collected in the vicinity of what is now Hong Kong (Appendix 2), while the remaining from either up the Pearl River towards Canton, or northwards along the Taiwan Strait at around 23°N. Of these Hong Kong records, 35 were new to science. Stimpson had planned for a comprehensive publication of the East Asian and Pacific crabs. To archieve this, he gathered the extensive collections of the NPEE, as well as the sizeable materials from the USEE reported by James Dwight Dana (1813–1895). Unfortunately, all this material was tragically lost in the Great Chicago Fire of 1871. Also lost were unpublished manuscripts and thousands of figures, some in color. Devastated, Stimpson died an early age of 40 in 1872, one year after the terrible fire. But not everything was lost. Three decades after Stimpson’s death, some manuscripts and illustrations of crustaceans, Brachyura and Anomura included, were recovered at the Navy Department at Virginia, partially edited by zoologist Mary Jane Rathbun (1860–1943) from the Smithsonian Institution, and were published posthumously as “Stimpson 1907”. Given the hostile atmosphere during the years of the American Civil War (1861–1865), in contrast with its predecessor, the USEE, which produced a good series of government documents, no official report was published for the NPEE (Deiss and Manning 1981; Vasile et al. 2005) (but see Habersham 1857; Cole 1947). The results of the NPEE, at least for the Brachyura, were very impressive when compared to similar scientific circumnavigations of similar scale in the 19th Century, such as the USEE (273 brachyuran species recorded, including 156 new species) (Dana 1852a b 1855) and the Challenger Expedition (305 species recorded with 49 new) (Miers 1886; Henderson 1888). The significance of Stimpson’s contributions was not only due to the sheer number of new taxa, among which many common species of shallow water and intertidal habitats; but also, as Raymond Manning commented, his “ability to accurately define so many taxa with only a few characters” (Vasile et al. 2005). In page 4 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan the intervening years, some of Stimpson’s specimens have been discovered in other museums, such as the U.S. National Museum of Natural History (Washington D.C.), the Natural History Museum (London) and the Museum of Comparative Zoology in Harvard University (AC Evans 1967; Griffin 1974; Deiss and Manning 1981; Manning and Reed 2006; Ng and Clark 2014; Ng 2017). These were specimens Stimpson had exchanged or given to colleagues before the fire. Unfortunately, the loss of most of his material has posed major challenges for subsequent generations of carcinologists. Through collections of topotypic material and/or selection of neotypes, however, many of Stimpson’s long synonymized taxa were resurrected, and the precise identities of others clarified (e.g., see Castro and Ng 2010; Shih et al. 2012; Komai and Poore 2016; Teng et al. 2016; Richer de Forges et al. 2021), testament to Stimpson’s genius. Shortly after the NPEE, an Austro-Hungarian vessel, S.M.S. Novara visited Hong Kong during their scientific circumnavigation from 1857 to 1859, staying in the vicinity of Hong Kong from 5th to 18th July 1858 (see narrative by Scherzer 1862), and the crustacean collections were later reported by Austrian zoologist Camill Heller (1823–1917) in 1862 and 1865. Eight species were reported from Hong Kong, all previously recorded by Stimpson (cf. Stimpson 1907). Two decades later, another circumnavigation, the United Kingdom’s Challenger Expedition (1873–1876), one first fully devoted to marine sciences (see narrative by Tizard et Fig. 3. Still-pristine out-lying islands of Hong Kong and Dangan Dao. A, B: aerial images showing current conditions. Extensive reclaimed land now as urban areas appearing light grayish white, in contrast to much darker vegetated areas. C: hand-drawn map drawn by Stimpson himself in the Expedition Journal. Sources: A, B: Google Earth, accessed Nov 2018; C: Vasile et al. (2005). page 5 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan al. 1885), stayed in vicinity of Hong Kong from 16th November 1874 to 8th January 1875, and surveyed a near-shore site near Hong Kong at a depth 7 to 10 fathoms. Five species of brachyurans were reported by zoologist Edward John Miers (1851–1930) in 1886, including a new record of Leucosia craniolaris (but see Galil 2003a, and accounts under List of Taxa below). German taxonomist Heinrich Otto Wilhelm Bürger (as O. Bürger; 1865–1945) reported on a small collection of sesarmid crabs collected by German biologist Carl Semper (1832–1893), en route to the Philippines in 1893 (Bürger 1893), and among the material were three species from Hong Kong. Hungarian traveler and scholar Count Béla Széchenyi (1837–1918) on his Eastern Asian Expedition (1877–1880) obtained a small collection of intertidal crustaceans while passing through Hong Kong and Macao. Eleven local records were presented in a report prepared by Hungarian zoologist Károly Koelbel in 1897 (Koelbel 1897), including one new species, the Buddhist crab Tmethypocoelis ceratophora (then as Dioxippe, Dotillidae), and three new records. During the U.S. Philippine Expedition (on the U.S.S. Albatross, 1907–1910), the vessel sailed northwards to Hong Kong for repair from early August to early November 1908 and surveyed a dozen of sites in “China Sea, vicinity Hongkong”. However, only several stations can be considered as within the territorial seas of Hong Kong (see definitions under Material and Methods below). Extensive collections and the growth of carcinology (early to mid20th Century) By the turn of the 20th century, as collections in major institutes increased, researchers focused on comparisons between similar forms across geographical regions. This fueled exchanges between relevant collections and new survey efforts globally. Based on material housed in the Indian Museum, Calcutta, among which included some from Hong Kong amassed by Surgeon-General Richard Hungerford (1834–1909), who travelled extensively in Asia (see mollusks reported by Nevill 1884), several India-based British zoologists referred to material from Hong Kong when reporting on the Indian fauna. In describing a raninid crab Lyreidus channeri Wood-Mason, 1885 (now Lysirude, Raninidae), entomologist James Wood-Mason (1846–1893), made comparisons, and named as well as illustrated a new species, Lyreidus stenops (Raninidae), from Hong Kong (Wood-Mason 1887). Physician and zoologist Alfred William Alcock (1859–1933) in his compilation of the Indian Brachyura (1895 1896 1898 1899 1900a b) included 24 species which had material from Hong Kong, contributing seven new local records. Marine biologist Stanley Wells Kemp (1882–1945) received a considerable lot of crustaceans, including various freshwater forms, amassed by zoologist Thomas Nelson Annandale (1876–1924), then Director of the Zoological Survey of India, collected on an excursion to the “Far East”: East and South China, to Thailand and Peninsular Malaysia. Two endemic freshwater decapods were recorded from Hong Kong, including a new species, Cryptopotamon anacoluthon (then as Potamon, Potamidae), from the Peak (Kemp 1918). In presenting zoological material from East Asia obtained by German zoologist Franz Theodor Doflein (1873–1924), German zoologist Heinrich Balss (1886– 1957) (see Balss 1922a b 1924) included some material from Hong Kong deposited in the collections of the Museum in Moscau, Russia (now Zoological Museum of Moscow University), in addition to several collected by German zoologist Hugo Hermann Schauinsland (1857–1937) during his excursion to East Asia around 1906 (see preface in Balss 1913). In these papers, 15 species from Hong Kong were reported, of which five were new records. Later in a revision of pilumnid crabs, Balss (1933) added Globopilumnus globosus (now Eupilumnus, Oziidae; see Ng et al. 2001) to the Hong Kong fauna based on material in the Museum of Kopenhagen (now the Zoological Museum, University of Copenhagen, Denmark). Swedish zoologist Nils Johan Teodor Odhner (as T. Odhner; 1879–1928), in reporting on xanthid crabs in 1925, reported four Hong Kong actaeine species from various institutes, of which Actaea ruppelli var. orientalis was described as new (now Gaillardiellus orientalis, Xanthidae). In 1926, American biologist Nathaniel Gist Gee (1876–1937), using past records and collections at the U.S. National Museum, Washington D.C., compiled a preliminary checklist of decapod crustaceans from China, including 98 brachyuran records from Hong Kong (or “near Hongkong”, “Hong Kong Islands” etc.), a figure corresponding to the report by Stimpson (1907). With substantial collections obtained by various naturalists, back at the British Museum (Natural History) (now the Natural History Museum), London, at the dawn of the 1930s, Scottish zoologist Isabella Gordon (1901–1988) published two important papers on brachyuran fauna of China which included Hong Kong (Gordon 1930 1931). The former paper (Gordon 1930) reported on a collection acquired by zoologist Richard William Durbin Barney, who was a Lecturer in Biology at the University of Hong Kong (HKU) until 1924. In this paper, five new portunid species were described, among which two, namely Charybdis (Goniosoma) barneyi and C. (Goniohellenus) sinensis have been considered as junior synonyms of C. affinis and C. vadorum, respectively (Yang et al. 2012; see Checklist page 6 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan below). Records enumerated in the latter paper (Gordon 1931) was based on another larger collection obtained by Barney, resulting in 95 entries, among which including one new portunid species, Neptunus (Hellenus) pulchricristatus, a replacement name for N. (H.) alcocki Gordon, 1930 which was preoccupied, as well as another 27 new records. Gordon (1934) also mentioned a peculiar antipatharian-associated Xenocarcinus tuberculatus White, 1847 (Epialtidae), which was a new and only record from Hong Kong when reporting on brachyuran material acquired from Indonesia. Hong Kong, as a part of the British Empire, had remained largely intact from military aggression until late 1941, and the preceding years had been fruitful for local brachyuran studies. This was substantially fueled by Chinese zoologist Chia-Jui Shen (also known as Jia-Rui Shen; 1902–1975). While investigating the brachyuran fauna of North China (Shen 1932b), he received considerable material amassed by British botanist and ornithologist Geoffrey Alton Craig Herklots (1902–1986), who was a Reader in Biology at HKU until 1928. This collection resulted in a series of four well-illustrated papers titled “Crabs of Hong Kong” (Shen 1931a b 1932a 1934), enumerating a total of 31 species, of which six were new records, with one new portunid species Charybdis (Goniohellenus) hongkongensis Shen, 1934 (now C. (Archias) hongkongensis). In the latter two papers (Shen 1932a 1934), a number of portunid crabs of substantial fishery values were treated. Shen himself had travelled to and collected in Hong Kong at least twice (1932 and 1938), and material he acquired resulted in two other papers, one in 1935 generally on taxa from coastal habitats, and the next was a checklist of Hong Kong brachyurans (Shen 1940a). The 1935 paper reported six local species, including new species Sesarma (Sesarma) gordoni (now considered as a junior synonym of Nanosesarma minutum, Sesarmidae) and Camptandrium anomalum (now Shenius), as well as one new record. The first checklist on Brachyura from Hong Kong by Shen (1940a) included past records and material deposited in National University of Peking, China, and the British Museum (Natural History), London, and 187 species were listed. This reported on 25 new records, including two new species, although the names were nomina nuda: Potamon (Potamon) hongkongensis (now Nanhaipotamon) and Cyclograpsus incisus (Varunidae). The two species were formally described in the later pages of the same issue [1(2)] of the Journal of the Hong Kong Fisheries Research Station (Shen 1940b); and two new records, Ilyoplax tenella (Dotillidae) and Gelasimus dussumieri (cited from Stimpson 1907 and collected from Canton River, and Gee 1926, collected from Macao, respectively). The latter probably represents the now common Tubuca paradussumieri (Ocypodidae). Post-WWII fisheries research (1950s–1980s) In December 1941, Hong Kong was occupied by Imperial Japan until the end of WWII in August 1945. During this period, studies of the local fauna came to a halt. Hong Kong’s population was 1,640,000 just before the Japanese occupation in 1941, but had plummeted to around 600,000 shortly after WWII. It subsequently soared from 1,860,000 in 1949, to 2,250,000 in the years after the Chinese Civil War and subsequent establishment of Communist China (see overview by SC Fan 1974). Endless political turmoil in mainland China forced Hong Kong to achieve self-sustainability in fisheries production in the post-war years (see Morton 2005). This had a major impact on marine discoveries. Wind-driven boats transitioned to mechanized fishing vessels after the war, expanding ranges and opportunities. The boats operated in many parts of Guangdong’s waters and reaching to eastern Hainan, at depths of about 110 m (Au 1968). Supporting fishery surveys were largely carried out at the Hong Kong Fisheries Research Station which was established in 1952. Much of the work was done in the South China Sea south of Hong Kong, with the surveys continuing into the 1980s (Chau and Wong 1960; Morton 2005). These surveys expanded our knowledge of several decapod groups (penaeid shrimps: Cheung 1960 1963, Wear and Stirling 1974; alpheid shrimps: Banner and Banner 1978; deep-sea lobsters: Bruce 1965a b 1966). For brachyurans, most remained unreported although samples have been included in other studies: Podophthalmus minabensis (now Vojmirophthalmus, Portunidae; Bruce 1965c), Lyreidus species (Raninidae, cf. Griffin 1970a), various spider crabs of the Majoidea (cf. Griffin and Tranter 1986), and more recently, a new majid species, Leptomithrax eldredgei (cf. Richer de Forges and Ng 2015). Several of these are apparently deeper-water forms and were probably collected from the edge or beyond the continental shelf from depths deeper than 100 m, and are unlikely to be from the shallower territorial waters of Hong Kong. Some of the material obtained from 1953 to 1984, is still probably extant and deposited in the Aberdeen Fisheries and Marine Offices, Hong Kong (see preliminary checklist by the Agricultural, Fisheries and Conservation Department (AFCD), https://www.hk-fish.net/english/ specimen_catalogue/introduction.html). This online list enumerated 73 brachyurans collected from 1958 to 1984, among which 58 of species-level identifications, and 43 have been confirmed in the literature or recent page 7 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan surveys (under List of Taxa below). A preliminary updated list of the brachyuran taxa is provided as appendix 3. These records, especially of deeper-water forms, need to be re-examined. Unfortunately, the first author did not manage to access the collection before departing for Taiwan in the Fall of 2020. Relevant records for this collection are excluded for the time being; the material should be re-examined in the future. Research workshops and international collaborations (1980s–2000s) Back in the 1930s when baseline data was generally absent, there was also a lack of experts in specific groups of local fauna. This was observed by British marine biologist Brian S. Morton (1942–2021) who joined HKU as a Lecturer in 1970, and embarked on a broad program to address the knowledge gap. In a career which spanned 33 years (retired in 2003), Morton mapped out strategies for wider surveys in Hong Kong and expanded collaborations between overseas and local researchers. From late 1970s onwards, Morton’s efforts resulted in many international workshops on marine biology, with invited experts from different fields and countries working in Hong Kong for varying periods of time, facilitating collaborations with local researchers and students on documentation of the local marine flora and fauna. At HKU, since the First International Workshop on Malacofauna of Hong Kong and Southern China, organized in 1977 (successively 1983 and 1992 on the same theme), subsequent events: “International Workshops on the Marine Flora and Fauna of Hong Kong and southern China” were held in 1980, 1986, 1989, 1995 and 1998. The earlier surveys were held at the YMCA Youth Camp at Wu Kai Sha, while those since early 1990s had been based at the Swire Institute of Marine Science (SWIMS) at Cape d’Aguilar, southeastern Hong Kong Island (see Morton 2003). Another one titled “International Workshop on Mangrove Ecosystem of Deep Bay and the Mai Po Marshes” in 1993 was organized jointly by the University and the World Wide Fund (WWF) for Nature, Hong Kong (SY Lee 1999). Themes of these events include identification and description of previously unknown taxa, identifying trends resultant to pollution, overfishing and other disturbances, with identification of species-rich areas for designation of conservation zones (Morton 2000 2003). Material were collected via numerous methods, and more notably, SCUBA diving was utilized for accessing subtidal fauna. The proceedings of the First International Biological Workshop organized in 1980 contained a number of key papers pertaining to the Brachyura: George (1982) on ghost crabs of the genus Ocypode (Ocypodidae), with description of one new species O. mortoni; Hill (1982) on leucosiid crabs; Bones (1982) on ecology of sentinel crabs Macrophthalmus (Macrophthalmidae); Dudgeon (1982) on decorating behavior of majid crabs; and Horikoshi and Takeda (1982) on assemblage of small subtidal crustaceans, particularly on some uncommon xanthoids. Several papers on parasitic isopods on decapod species (Markham 1982 1990 1992; summarized by Li 2003) were also part of the proceedings, and their hosts included two new brachyuran records. Some of these results, as well as those of the Flora and Fauna checklists (Hill et al. 1975 1978), were incorporated into The Sea Shore Ecology of Hong Kong (Morton and Morton 1983), which became a must-read for students of Hong Kong marine biology. Subsequent workshops added papers on the brachyuran fauna: Pregenzer and Morton (1990) on six new records of bivalve-commensal pinnotherid crabs; Davie (1992a) on intertidal brachyuran fauna with one new species Indopinnixa mortoni (Pinnotheridae) and five new records; Davie (1992b) on subtidal assemblages of anomuran and brachyuran crabs in Tolo Harbour, Tolo Channel and Mirs Bay derived from trawl surveys, with two new records; and SY Lee and Leung (1999) contributed five new records and life photographs of intertidal crabs, particularly sesarmids from Mai Po and Deep Bay. Investigating epibenthic brachyurans on southeastern seas of Hong Kong and changes of assemblages as result of extensive trawling and dredging during the 1990s, Blackmore and Rainbow (2000) and SY Lee et al. (2000) contributed more subtidal records, with Ng and Dai (1997) describing a new pilumnid genus and species, Xestopilumnus cultripollex, from these samples. In 2002, the Hong Kong Submarine Caves Expedition, jointly organized by SWIMS and the Natural History Museum (London), collected samples from two marine caves at Conic Island and Steep Island (Morton and Bamber 2008). Results of these surveys on decapods were presented by Clark et al. (2008). The six brachyuran reported included one new pilumnid species, Latopilumnus conicus (described by Ng and Clark 2008 in the same volume) and three new records. Detailed reports, field guides, and recent progress (1970s onwards) Numerous papers on various groups of Brachyura were published in the late 1970s and 1980s, some treating the taxa in considerable depth and detail. American zoologist Jocelyn Crane (1909–1998) visited Hong Kong for several days in 1963, investigating page 8 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan taxonomy, behavior and evolution of fiddler crabs, eventually presenting in her seminal work Fiddler Crabs of the World. She collected in the vicinity of the Castle Peak area, and five species of fiddler crabs were collected (Crane 1975). Of this material, one Uca (Thalassuca) vocans borealis (now Gelasimus borealis; see Shih et al. 2015) was described as new, and a neotype of Tubuca acuta was also designated. Singaporean biologist Cheng Lam Soh visited and collected in Hong Kong in 1975, and in 1978, published a report on 15 sesarmid crabs, with description of three new species, namely Chiromantes maipoensis (now Parasesarma; see Shahdadi and Schubart 2017), Pseudosesarma patshuni (now Orisarma patshuni; see Schubart and Ng 2020) and Holometopus serenei (a junior synonym of Chiromantes haematocheir), and added two new records. Studying intertidal sipunculanassociated fauna, American zoologist Raymond Brendan Manning (1934–2000), in collaboration with Brian Morton, described one new pinnotherid genus and species, Indopinnixa sipunculana (Manning and Morton 1987). Singaporean zoologist Peter K. L. Ng visited Hong Kong in 1991, collaborated with freshwater biologist David Dudgeon at HKU, and described a new gecarcinucid species Somanniathelphusa zanklon based local material (previously collectively under the name S. sinensis (H. Milne Edwards, 1853); see below), as well revised the taxonomic status of two other endemic freshwater crabs (Ng and Dudgeon 1992). Some of this material (deposited in Singapore) was subsequently examined and reported by Chinese zoologist Ai-Yun Dai (1930–2005) and used in her monumental monograph on Parathelphusidae and Potamidae in China under the series Fauna Sinica (Dai 1999) (see also Ng 2000a). After the defining work on fiddler crabs (then collectively as Uca) by Crane (1975), Australian zoologist Diana S. Jones, in collaboration with Brian Morton, reported and illustrated six local species in detail (Jones and Morton 1994). Fiddler crab taxonomy, including again local and southern Chinese taxa, were revised by Taiwanese zoologist Hsi-Te Shih and colleagues (2010b), and later a review of the ocypodid species Uca crassipes (Shih et al. 2012), in which Gelasimus splendidus, described based on material from Hong Kong and long considered synonymous with the former, was resurrected. In the 2000s, a number of important guidebooks of fauna from various habitats were published. Although none were solely on brachyuran crabs, nor any new records presented, many of those were well-illustrated and user-friendly, providing an invaluable resource for public education. One of the earliest of this kind was a beautiful A Colour Guide to Hong Kong Animals published in 1981 by Dennis S. Hill, senior lecturer of zoology, HKU, in collaboration with illustrator Karen Phillipps (the Chinese version published in 1982). This guidebook presented 20 common brachyuran species illustrated in color and compelling detail. On coastal and shallow marine habitats, Brian Morton, in collaboration with various authors, published several other guidebooks or pamphlets introducing specific sites: Mai Po (Melville and Morton 1983), Hoi Ha Wan (Morton and Ruxton 1992 1997) and Cape d’Aguilar (Morton and Harper 1995). On mangrove habitats, S. Y. Lee published Venturing Forests in the Water in 2003 (SY Lee 2003). Several series of guidebooks had further included the local brachyuran fauna: Hong Kong Mangroves by Nora F. Y. Tam and Y. S. Wong at the City University of Hong Kong in 2000, three (four in the series) in 2003 by biologists at HKU, namely Rocky Shores by Gray A. Williams, Hillstreams by David Dudgeon, and Sandy Shores by Benny K. K. Chan and Kevin J. Caley, followed by Commercial Crustaceans in Hong Kong in 2004 by Siu Fai Leung and colleagues at the Agricultural, Fisheries and Conservation Department, HKSAR Government, and later another series by the team of local zoologists Terrence C. W. Fong, Vincent C. S. Lai, Henry T. H. Lui and Samson N. H. So (Estuarine Organisms (2005), Hard Shore Organisms (2006) and Small Wetland Creatures (2007)). Our understanding of the local brachyuran fauna has been a slow but steady process. Vincent Lai, in 1999, reported two new intertidal records, namely Ebalia malefactrix and Episesarma versicolor (Leucosiidae and Sesarmidae, respectively), in Porcupine!, the newsletter of the Department of Ecology and Biodiversity, HKU (VCS Lai 1999a b). In establishing a new subfamily Gaeticinae under the Varunidae, Peter Davie and Singaporean zoologist Ngan Kee Ng (Davie and NK Ng 2007) reported on three local species under the group, and that of Gaetice ungulatus Sakai, 1939 was new. Similarly, the revision of the Parasesarma plicatum species-group (Rahayu and Ng 2010), resurrected P. affine (De Haan, 1837) (Sesarmidae) in East Asia as a valid species and added a new local record of P. ungulatum (H. Milne Edwards, 1853). Noteworthy is Leptomithrax eldredgei Richer de Forges & Ng, 2015 (Majidae), described from material collected in surveys by the Hong Kong Fisheries Research Station back in the 1960s (see above). The discovery of an intriguing tree-climbing sesarmid crab Haberma tingkok Cannicci & Ng, 2017, found exclusively among canopy of mangroves, was also a significant one as it has substantial conservation significance. In investigating small scavenging swimming crabs of the genus Eodemus (previously Portunus or Xiphonectes), local zoologist Kingsley J. H. Wong, in collaboration with colleagues in Hong Kong and Taiwan, added one page 9 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan to 2000s; and (4) additions by naturalists throughout the recent decade. A multi-decadal hiatus can be observed from 1950s to 1970s after the WWII. As such, the history of species inventory has been shaped by contributions from series of work by carcinologists, as well as the prevailing social-economical circumstances. As to address whether the local brachyuran fauna being anyway well documented, the best answer is “appears to be, but far from being well understood”. Apart from as indicated from the steep, upward cumulative curve (Fig. 5A), even for intertidal habitats which are more accessible to naturalists and researchers alike, there are no short of surprises—the long-legged arboreal Haberma tingkok (Sesarmidae) was only recently discovered among dense mangrove canopies (Cannicci and Ng 2017). Shallow rocky reefs and coral communities still harbor many unrevealed taxa (see below). Noteworthy is that there are still no reliable reports of any members of the Superfamily Trapezioidea from Hong Kong, other than a questionable record of Tetralia glaberrima (Herbst, 1790) (Tetraliidae, see below). No members of the widespread families Trapeziidae or Domeciidae, all of which are obligate symbionts of hard corals, have been reported thus far, although scleractinians are present in Hong Kong. Similarly, for cryptochirids, obligate symbionts of scleractinians, the genetic diversity of this group has been grossly underestimated in previous decades. In addition, through revisionary works adopting morphological and molecular approaches, cryptic forms continue to be identified, establishing new and elucidating previously synonymized taxa (e.g., Wong et al. 2010a; Shih et al. 2012; Nguyen and Ng 2021). We hope this paper presents a comprehensive and accurate summary of the brachyuran species so far known from Hong Kong, and will facilitate future investigations on the fauna, and associated studies on ecology, biology and conservation. MATERIALS AND METHODS Recognizing records from Hong Kong The location of Hong Kong along the coasts of South China, and a map of Hong Kong is shown in Figure 1. Given the multi-staged evolution of the understanding of the locality of Hong Kong outlined under “Historical Account” above, older pre-20th century records as “near Hong Kong” or “off Hong Kong” are accepted as local records (Fig. 2). Another complication comes from various records from deeper water, or species normally collected from seas of the continental shelf in northern South China Sea. While the territorial seas of Hong Kong are generally less than 50 m of depth (e.g., see Fyfe et al. 2000 for bathymetry map), reports derived from localities apparently outside this range, such as two new species of euxanthine crabs reported from “South China Sea, near Hong Kong”, from depths of 180 and 203 m (Ng and Chen 2005), which are not in the now recognized vicinity, are thus excluded. For records with GPS (Global Positioning System) data, an arbitrary limit is taken as followings, as to exclude those surveyed significantly distant from Hong Kong: southern limit at 21°N, and western and eastern limits along 113° and 115°E, respectively. This approach leaves a “working zone” of around 100 km in radius at the northern portion of the South China Sea. Records from outside of this zone are excluded. Older records indicated such as “near Hong Kong”, or “South China Sea near Hong Kong” without depth, precise location data or GPS, have to be accepted for the sake of completeness. Selection of formal records There is a great deal of literature on crabs, not just scientific reports, degree theses, guide books, pamphlets and other educational material, but also newsletters, newspaper articles, aquarist writings, and recipes of all kinds, and in the recent decades, an explosion of online blogs, social media, databases and so forth. Compiling an exhaustive checklist including all of these sources is impractical, so we have to be more targeted. We treat publications as defined in the International Code of Zoological Nomenclature (Article 8 in ICZN 2012)— regardless in printed or electronic forms, in general, one which considered publicly disseminated, with clear indications of date of publication, obtainable, and presenting unamendable scientific information. We agree with these criteria as the data captured should be accurate. The online database Hong Kong Register of Marine Species (HKRMS), itself a subset of World Register of Marine Species (WoRMS), includes online entries which are amendable over time, and contains many unpublished works like various local degree dissertations, government and consultant reports. These documents are generally not available to the public; the data are not readily obtainable or verifiable by researchers and are essentially unrefereed. As such, we prefer to omit entries now listed on HKRMS. Similarly, degree theses and dissertations are not readily accessed by most workers, and results presented are considered unpublished data by respective authors, thus relevant records are herein excluded. In this work, emphasis is placed on references containing information on crab morphology, taxonomy, diversity, and distribution; and pertaining to Hong Kong as far as possible. As page 16 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan for the multitude of papers published on brachyurans that treat ecology, physiology and other biological aspects etc., they are generally excluded unless new records and/or new taxonomic ideas are presented (e.g., Dudgeon 1982). There are some in which the original identifications are dubious or likely erroneous, and we have amended them. These are justified under Remarks of respective entries. Overall, we are confident that this layered approach strikes a balance between presenting a comprehensive checklist of fauna and keeping the data accurate and manageable. Format of catalogue The classification scheme of the present catalogue follows that of Davie et al. (2015a b) which is revised and updated from Ng et al. (2008). Exceptions, such as classifications of the Portunoidea updated recently by Spiridonov et al. (2014), N. Evans (2018), Spiridonov (2020) and Koch et al. (2022), as well as that of the Sesarmidae by Schubart and Ng (2020), will be stated under respective Remarks. Under each species entry consists of the following information. (1) Valid scientific name and authority: full citations of species authority are not provided for simplicity. Readers are suggested to consult works listed under “Taxonomy” (see below). (2) Citation of local works of the species are arranged in chronological order. In bilingual work only the English portion would be cited. In works published separately in two languages, between which barely differs (e.g., Hill and Phillipps (1981) in English, and Hill and Phillipps (1982) in Chinese), only the English version would be included. As for those with versions substantially modified (e.g., Morton and Ruxton (1992) in English; Morton and Ruxton (1997), bilingual version), both are treated as separate works. For first and sole authors with shared family names (particular of Asian ethnic origins), the initials also are indicated (e.g., CGS Tan and SH Tan), in exception of H-L Chen, PKL Ng, T Sakai, H-T Shih and KJH Wong, who, at least so far, published or recorded more local taxa than, respectively, W-J Chen, NK Ng, K Sakai, Y-J Shih and ECK Wong. (3) Taxonomy: important references providing taxonomic information fostering precise identifications, but not meant to be exhaustive. (4) Type: whereabouts and status of type material. For extant material, acronym of institute deposited (full names listed in Appendix 4), and access number of name-bearing type, if applicable, is indicated. (5) Type locality: current name of locality indicated if possible. (6) Distribution; and (7) Habitat, with scientific names and authority of hosts of symbiotic brachyurans (cryptochirids and pinnotherids) following current formats indicated on WoRMS. (8) Remarks: additional taxonomic information justifying use of names and identity of local records. Throughout the past decade (2010s), the first author (KJHW) had access to brachyuran material largely from benthic trawl surveys (reported in Wong et al. 2021); routine shore surveys; among fouling communities; and habitats in vicinity of coral communities by scuba diving. Although this paper presents results of a largely “desk study” approach, and the material encountered in this time period cannot be seen as an exhaustive representation of the fauna, it would be useful to indicate the taxa which local material was examined in this timespan. This indicated with a superscript hash symbol (#) after the title of each species entry, following the authority of species. New records presented are provided with illustrations showing diagnostic features including photographs and line drawings. Images showing overall habitus of preserved specimens are indicated as monochrome colorations in figures 35 to 41. The morphological terminology follows that of Davie et al. (2015c). Material of most of the illustrated taxa are deposited at the Coastal Ecology Laboratory (CEL), Academia Sinica, Taipei, the Zoological Collections of the Department of Life Science, National Chung Hsing University (NCHUZOOL), Taichung, Taiwan; and the Zoological Reference Collection (ZRC) of the Lee Kong Chian Natural History Museum, National University of Singapore, Singapore. Depository of illustrated specimens is listed in appendix 5. Ongoing research on several intertidal forms under preparation by us and colleagues are briefly elaborated under respective Remarks, and preliminarily included in the present work for completeness. LIST OF SUPERFAMILIES, FAMILIES AND SUBFAMILIES OF BRACHYURAN CRABS RECORDED FROM HONG KONG Number of species in each superfamily are indicated in curly brackets ({ }), and that of lowest supra-generic groupings (family or subfamily level) in square brackets ([ ]). INFRAORDER BRACHYURA LATREILLE, 1802 SECTION PODOTREMATA GUINOT, 1977 Superfamily DROMIOIDEA De Haan, 1833 {4} Family Dromiidae De Haan, 1833 Subfamily Dromiinae De Haan, 1833 [3] Family Dynomenidae Ortmann, 1892 Subfamily Dynomeninae Ortmann, 1892 [1] Superfamily RANINOIDEA De Haan, 1839 {3} Family Lyreididae Guinot, 1993 [2] Family Raninidae De Haan, 1839 [1] page 17 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan SECTION EUBRACHYURA SAINT LAURENT, 1980 SUBSECTION HETEROTREMATA GUINOT, 1977 Superfamily AETHROIDEA Dana, 1851 {1} Family Aethridae Dana, 1851 [1] Superfamily CALAPPOIDEA De Haan, 1833 {9} Family Calappidae De Haan, 1833 [7] Family Matutidae De Haan, 1835 [2] Superfamily CARPILIOIDEA Ortmann, 1893 {1} Family Carpiliidae Ortmann, 1893 [1] Superfamily CORYSTOIDEA Samouelle, 1819 {2} Family Corystidae Samouelle, 1819 [2] Superfamily DAIROIDEA Serène, 1965 {1} Family Dairidae Serène, 1965 [1] Superfamily DORIPPOIDEA MacLeay, 1838 {7} Family Dorippidae MacLeay, 1838 [7] Superfamily ERIPHIOIDEA MacLeay, 1838 {7} Family Eriphidae MacLeay, 1838 [2] Family Menippidae Ortmann, 1893 [2] Family Oziidae Dana, 1851 [3] Superfamily GECARCINUCOIDEA Rathbun, 1904 {1} Family Gecarcinucidae Rathbun, 1904 [1] Superfamily GONEPLACOIDEA MacLeay, 1838 {12} Family Euryplacidae Stimpson, 1871 [7] Family Goneplacidae MacLeay, 1838 Subfamily Goneplacinae MacLeay, 1838 [4] Family Scalopidiidae Števčić, 2005 [1] Superfamily HEXAPODOIDEA Miers, 1886 {4} Family Hexapodidae Miers, 1886 [4] Superfamily HYMENOSOMATOIDEA MacLeay, 1838 {3} Family Hymenosomatidae MacLeay, 1838 Subfamily Hymenosomatinae MacLeay, 1838 [3] Superfamily LEUCOSIOIDEA Samouelle, 1819 {33} Family Iphiculidae Alcock, 1896 [2] Family Leucosiidae Samouelle, 1819 Subfamily Cryptocneminae Stimpson, 1907 [1] Subfamily Ebaliinae Stimpson, 1871 [23] Subfamily Leucosiinae Samouelle, 1819 [7] Superfamily MAJOIDEA Samouelle, 1819 {26} Family Epialtidae MacLeay, 1838 Subfamily Epialtinae MacLeay, 1838 [5] Subfamily Pisinae Dana, 1851 [10] Family Inachidae MacLeay, 1838 Subfamily Inachinae MacLeay, 1838 [4] Family Majidae Samouelle, 1819 Subfamily Majinae Samouelle, 1819 [7] Superfamily ORITHYIOIDEA Dana, 1852 {1} Family Orithyiidae Dana, 1852 [1] Superfamily PALICOIDEA Bouvier, 1898 {1} Family Palicidae Bouvier, 1898 [1] Superfamily PARTHENOPOIDEA MacLeay, 1838 {7} Family Parthenopidae MacLeay, 1838 Subfamily Parthenopinae MacLeay, 1838 [7] Superfamily PILUMNOIDEA Samouelle, 1819 {43} Family Galenidae Alcock, 1898 Subfamily Dentoxanthinae Števčić, 2005 [1] Subfamily Galeninae Alcock, 1898 [1] Subfamily Halimedinae Alcock, 1898 [2] Family Pilumnidae Samouelle, 1819 Subfamily Eumedoninae Dana, 1852 [4] Subfamily Pilumninae Samouelle, 1819 [26] Subfamily Rhizopinae Stimpson, 1858 [9] Superfamily PORTUNOIDEA Rafinesque, 1815 {49} Family Carcinidae MacLeay, 1838 Subfamily Polybiinae Ortmann, 1893 [1] Family Geryonidae Colosi, 1924 Subfamily Ovalipinae Spiridonov, Nerelina & Schepetoc, 2014 [1] Family Portunidae Rafinesque, 1815 Subfamily Lupocyclinae Paulson, 1875 [2] Subfamily Necronectinae Glaessner, 1928 [3] Subfamily Podophthalminae Dana, 1851 [1] Subfamily Portuninae Rafinesque, 1815 [11] Subfamily Thalamitinae Paulson, 1875 [30] Superfamily POTAMOIDEA Ortmann, 1896 {3} Family Potamonidae Ortmann, 1896 Subfamily Potamiscinae Bott, 1970 [3] Superfamily TRAPEZIOIDEA Miers, 1886 {1} Family Tetraliidae Castro, Ng & Ahyong, 2004 [1] Superfamily XANTHOIDEA MacLeay, 1838 {39} Family Xanthidae MacLeay, 1838 Subfamily Actaeinae Alcock, 1898 [9] Subfamily Banareiinae Števčić, 2005 [1] Subfamily Chlorodiellinae Ng & Holthuis, 2007 [3] Subfamily Cymoinae Alcock, 1898 [1] Subfamily Etisinae Ortmann, 1893 [2] Subfamily Euxanthinae Alcock, 1898 [2] Subfamily Kraussiinae Ng, 1993 [1] Subfamily Liomerinae Sakai, 1976 [4] Subfamily Xanthinae MacLeay, 1838 [9] Subfamily Zosiminae Alcock, 1898 [7] SUBSECTION THORACOTREMATA GUINOT, 1977 Superfamily CRYPTOCHIROIDEA Paul’son, 1875 {7} Family Cryptochiridae Paul’son, 1875 [7] Superfamily GRAPSOIDEA MacLeay, 1838 {51} Family Grapsidae MacLeay, 1838 [3] Family Percnidae Števčić, 2005 [1] Family Plagusiidae Dana, 1851 [2] Family Sesarmidae Dana, 1851 [22] Family Varunidae H. Milne Edwards, 1853 Subfamily Asthenognathinae Stimpson, 1858 [2] Subfamily Cyclograpsinae H. Milne Edwards, 1853 [9] Subfamily Gaeticinae Davie & N. K. Ng, 2007 [3] Subfamily Varuninae H. Milne Edwards, 1853 [9] Superfamily OCYPODOIDEA Rafinesque, 1815 {49} Family Camptandriidae Stimpson, 1858 [7] Family Dotillidae Stimpson, 1858 Subfamily Dotillinae Stimpson, 1858 [12] Subfamily Sheniinae Ng, Clark & Cuesta, 2010 [1] Family Macrophthalmidae Dana, 1851 Subfamily Ilyograpsinae Števčić, 2005 [1] Subfamily Macrophthalminae Dana, 1851 [10] Subfamily Tritodynamiinae Števčić, 2005 [3] Family Mictyridae Dana, 1851 [1] Family Ocypodidae Rafinesque, 1815 Subfamily Gelasiminae Miers, 1886 [8] Subfamily Ocypodinae Rafinesque, 1815 [3] Family Xenophthalmidae Stimpson, 1858 Subfamily Anomalifrontinae Rathbun, 1931 [1] Subfamily Xenophthalminae Stimpson, 1858 [2] page 18 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Superfamily PINNOTHEROIDEA De Haan, 1833 {17} Family Pinnotheridae De Haan, 1833 Subfamily Pinnothereliinae Alcock, 1900 [4] Subfamily Pinnotherinae De Haan, 1833 [13] LIST OF TAXA Section Podotremata Guinot, 1977 Superfamily Dromioidea De Haan, 1833 Family Dromiidae De Haan, 1833 Subfamily Dromiinae De Haan, 1833 Conchoecetes atlas McLay & Naruse, 2019 # Conchœcetes artificiosus — Stimpson 1858d: 240; Alcock 1900a: 151; Stimpson 1907: 180, pl. 21(5) [not Dromia artificiosus Fabricius, 1798]. Conchoecetes artificiosus — Gee 1926: 160 [list; “off Hong Kong”]; Davie 1992b: tab. 1; Ng et al. 2000: 156, fig. 1a, b [not D. artificiosus Fabricius, 1798]. Conchocoetes artificiosus — Morton and Morton 1983: 95 [not D. artificiosus Fabricius, 1798]. Conchoecetes atlas McLay & Naruse, 2019: 10, figs. 4, 5, 6B; Wong et al. 2021: 6, fig. 2, pl. 1A. Taxonomy: Ng et al. (2000: as C. artificiosus), McLay and Naruse (2019). Type: Holotype ♂ ZRC 2019.1631. Type locality: Camiguin Island, Bohol Sea, the Philippines. Distribution: West Pacific: Indonesia, Singapore, Philippines, Gulf of Thailand, South China, Taiwan and Japan (McLay and Naruse 2019). Habitat: Subtidal, muddy or sandy substrates; depths 30 to 150 m (Chen and Sun 2002). Remarks: Past local records of “C. artificiosus” probably represent C. atlas or C. chanty (see McLay and Naruse 2019). On the basis of material on hand (reported by Wong et al. 2021), characterized by a sub-pentagonal carapace, bearing a distinct obtuse anterolateral tooth but devoid of posterolateral tooth (Wong et al. 2021: fig. 2a, pl. 1A), we can record C. atlas McLay & Naruse, 2019 for now. Lauridromia dehaani (Rathbun, 1923) # Dromia rumphii — Stimpson 1858d: 240; Stimpson 1907: 177, pl. 21(7). [not Dromia rumphii Fabricius, 1798 = Cancer dormia Linnaeus, 1763]. Dromia dehaani Rathbun, 1923: 68; Gee 1926: 160; Gordon 1931: 526; Morton 1988: 22, pl. 3; Davie 1992b: tab. 1; Blackmore and Rainbow 2000: app. 1. Dromia dormia — Shen 1931a: 96, text-figs. 3, 4, pl. 4; Hill et al. 1975: 65; Hill et al. 1978: 92; Morton 1979a: 121, fig. 6.19; Morton 1979b: 129; Hill and Phillipps 1981: 156, pl. 52i; Morton and Morton 1983: 95, fig. 10.4(8) [not Cancer dormia Linnaeus, 1763]. Lauridromia dehaani — Dai and Ng 1997: 754, fig. 1; Sin et al. 2009: tab. 2; Wong et al. 2021: 7, fig. 3, pl. 1B. Taxonomy: Rathbun (1923: as Dromia), McLay (1993), Ahyong et al. (2009). Type: Syntypes in USNM, RMNH, NHM (Yamaguchi and Baba 1993; Fransen et al. 1997). Type locality: Hakodate, Japan. Distribution: Western Indian Ocean to West Pacific: from South and East Africa, Red Sea, Pakistan, India, to Indonesia, South and East China Seas and East Asia. Habitat: Subtidal, muddy substrates; depths 14 to 150 m (Chen and Sun 2002); occasionally intertidal. Remarks: Rathbun (1923) gave a thorough elaboration on the distinctions of two large-sized dromiids in the Indo-Pacific, at present represented by the names Tumidodromia dormia (Linnaeus, 1763) and Lauridromia dehaani (Rathbun, 1923). The name Dromia rumphii Fabricius, 1798, was shown to be a junior synonym of Cancer dormia Linnaeus, 1763. However, this name was used by numerous earlier authors to represent a form found East Asia, including De Haan (1839: material from Japan) and Stimpson (1858d: material from Hong Kong). This form was recognized as D. dehaani by Rathbun (1923). The records of “D. dormia” from Hong Kong were based on an illustrated but erroneous record in Shen (1931a) (Rathbun 1923; Sakai 1976; Dai and Yang 1991; Chen and Sun 2002). Petalomera granulata Stimpson, 1858 Petalomera granulata — Gordon 1931: 526; Blackmore and Rainbow 2000: app. 1. Taxonomy: Sakai (1936), McLay and Ng (2007). Type: Neotype ♀ SMF (designated by McLay and Ng 2007). Type locality: Sagami Bay, Japan (neotype). Distribution: East Asia: South China Sea, Taiwan and Japan (Chen and Sun 2002). Habitat: Subtidal, substrates sandy, muddy, or shelly; depths of 50 to 150 m (Chen and Sun 2002). Family Dynomenidae Ortmann, 1892 Subfamily Dynomeninae Ortmann, 1892 Hirsutodynomene spinosa (Rathbun, 1911) Dynomene cf. spinosa — Blackmore and Rainbow 2000: app. 1. Taxonomy: McLay (1999), Chen and Sun (2002), Ahyong et al. (2009). Type: Holotype ♂ USNM 41048 (McLay 1999). page 19 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Type locality: Coetivy Island, Seychelles. Distribution: Western Indian Ocean to Central Pacific: from Madagascar to Indonesia, Southeast and East Asia, Australia, Mariana Islands, Guam, Line Islands and French Polynesia (McLay 1999; Chen and Sun 2002). Habitat: Intertidal; shallow coral reeds; depths up to 15 m (McLay 1999). Superfamily Raninoidea De Haan, 1839 Family Lyreididae Guinot, 1993 Lyreidus stenops Wood-Mason, 1887 Lyreidus stenops Wood-Mason, 1887: 209, pl. 1(7, 8); Griffin 1970a: 106, figs. 1, 6c, h, q, 7e, 8d, pl. 2A [south of Hong Kong: 21°10.7'N to 21°20'N, 114°25.6'E to 114°28.5'E]. Lyreidus politus — Gordon 1931: 532; Shen 1940a: 214. Lyreidus integra — Shen 1931b: 190, text-fig. 7, pl. 12(2). Taxonomy: Griffin (1970a), Goeke (1985), Ahyong et al. (2009). Type: Holotype ♀ IM-8467/6 (Griffin 1970a). Type locality: Hong Kong. Distribution: Eastern Indian Ocean to West Pacific: Western Australia, South and East China Seas, Taiwan and Japan (Chen and Sun 2002; Ahyong et al. 2009). Habitat: Subtidal; substrates sandy; depths 31 to 118 m (Ahyong et al. 2009). Lyreidus tridentatus De Haan, 1841 Lyreidus 3-dentatus — Wood-Mason 1887: 209. Lyreidus tridentatus — Griffin 1970a: 94, figs. 1–5, 6a, e, f, j–o, 7a, b, 8a, b, f, g, pl. 1A [south of Hong Kong: 21°04.5'N to 21°15.2'N, 114°30.5'E to 115°18'E]. Taxonomy: Griffin (1970a), Goeke (1985), Ahyong et al. (2009). Type: Lectotype ♀ RMNH D 23017 and paralectotypes in RMNH (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: West to Central Pacific: East Asia to Australia, New Zealand, New Caledonia, Fiji and Hawaii (Chen and Sun 2002; Ahyong et al. 2009). Habitat: Subtidal; substrates sandy-mud shelf on shelf and slope; depths 27 to 382 m (McLay 1988). Family Raninidae De Haan, 1839 Ranina ranina (Linnaeus, 1758) Ranina ranina — Shen 1931a: 103, text-fig. 9, pl. 7; Shen 1940a: 214. Taxonomy: Ahyong et al. (2009). Type: Status unknown, probably lost (Davie 2002). Type locality: Indian Ocean. Distribution: Western Indian Ocean to Central Pacific: South and East Africa to Australia, East Asia, Hawaii and French Polynesia (Chen and Sun 2002; Ahyong et al. 2009). Habitat: Intertidal to shallow subtidal; substrates sandy, to depths of 121 m (Ahyong et al. 2009). Section Eubrachyura Saint Laurent, 1980 Subsection Heterotremata Guinot, 1977 Superfamily Aethroidea Dana, 1851 Family Aethridae Dana, 1851 Drachiella morum (Alcock, 1896) # Drachiella morum — Wong et al. 2021: 7, fig. 4a, b, pl. 1C. Taxonomy: Serène (1954: as Actaeomorpha), Serène and Soh (1976), Chen (1989). Type: Syntype ♀♀ in ZSI (Chopra 1934). Type locality: Ganjam coast, northeastern India, Gulf of Bengal. Distribution: Eastern Indian Ocean to South China Sea and East Asia: Bay of Bengal, Phuket (Thailand), South and East China Seas, and Japan (Chen and Sun 2002). Habitat: Subtidal: substrates of muddy sand, coarse silt or shell fragments; depths 23 to 150 m (Chen and Sun 2002). Superfamily Calappoidea De Haan, 1833 Family Calappidae De Haan, 1833 Calappa calappa (Linnaeus, 1758) Calappa calappa — Hill et al. 1978: 114; Morton 1979b: 136; Morton and Morton 1983: 158, fig. 10.4(5). Taxonomy: Chen (1993), Galil (1997), Chen and Sun (2002). Type: Lost (see discussion below). Type locality: Asia. Distribution: Western Indian Ocean to Central Pacific: from East Africa, Madagascar, India, to Indonesia and East Asia, and New Caledonia, French Polynesia and Hawaii (Galil 1997; Chen and Sun 2002). Habitat: Subtidal: rocky, gravelly and shelly bottoms, of depth 10 to 50 m (Chen and Sun 2002). Remarks: Calappa calappa was described by Linnaeus (1758) from somewhere in Asia. Cancer fornicatus Fabricius, 1781, is generally regarded as a junior synonym, although the types of both nominal page 20 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan taxa have never been redescribed, figured or reexamined since their original descriptions. Galil (1997) listed in her synonymy two other species: Cancer flosculosus Seba, 1759, and Cancer heracleoticus Seba, 1759, but without any comments. Linnaeus (1758: 630) described Cancer calappa very briefly, “C. brachyurus, thorace strigus fubimbricato gibbo trilobo.” He did not indicate if he had any specimens with him, but he did refer to Rumphius (1705: 21, pl. 11 figs. 2, 3) record of this species from Ambon (Indonesia). The type locality was stated as from somewhere in Asia. In the Stockholm Museum in Uppsala, Sweden (UMS), where Linnaeus’ material is now kept, there is specimen of Cancer calappa Linnaeus, 1758 (see Holm 1957: 56). The specimen was examined and thoroughly photographed through the courtesy of S.H. Tan who visited the museum at the request of the second author. The specimen (UMS 254), a dried male measuring 70.8 by 125.2 mm, is still in good condition. This specimen was labelled as from the collection of “Gustav IV Adolf”, which was obtained by Linnaeus only after 1758 and subsequently reported in 1764 (see Holm 1957: 56; Wallin 1997: 16). This specimen, however, is not one of Linnaeus’ type(s) of Cancer calappa. Specimen UMS 254 was obtained only after 1758 and therefore cannot be a type. Linnaeus’ type specimen(s) of Cancer calappa Linnaeus, 1758 (if indeed he had any), is in all probability, lost. As Linnaeus (1758) also described C. calappa with reference to Rumphius’ (1705) work, Rumphius’ material could also constitute the type series. Unfortunately, the latter’s specimens are also lost. The type specimen of Cancer fornicata Fabricius, 1781, a dried male in the Natural History Museum of Denmark in the University of Copenhagen, is clearly a subjective junior synonym of Cancer calappa Linnaeus, 1758. Fabricius (1781) did not indicate how many specimens he had with him, and as such, the only extant specimen is regarded as the lectotype of C. fornicata. The identities of Seba’s (1759) Cancer flosculosus and C. heracleoticus is more problematic. The two taxa in question were published in the third volume of the series which is often cited as “1758” or “1761”, but Holthuis (1969) showed that it was released in 1759. In any case, Seba’s (1759) names are not available for nomenclatural purposes. The beginning of the Seba’s (1759) book (in the contents section after the preface) has an unpaginated summary list of the plates, with an associated index (in Latin) and table (in French), and this has names which appear to be binominal scientific names. Subsequently, Seba appended a similar Latin index and French table after each plate to elaborate on the figures, and it often contains what appear to be binomial names in Latin. The problem with Seba’s (1759) work is that in his Latin indices, he uses a mixture of binomial as well as polynomial names throughout the book. For the Decapod Crustacea, while most of these “scientific names” are in Latin and appear binomial, he also uses names like “Cancer pagurus Maja” (for numbers 2 and 3 of his plate 18), “Cancer Uka una” (for number 8 of his plate 18), and “Cancer Ciri Apoa” (for number 9 of plate 18). Article 11.4 states that for names to be nomenclaturally available, there must be “Consistent application of binominal nomenclature. The author must have consistently applied the Principle of Binominal Nomenclature [Art. 5.1] in the work in which the name or nomenclatural act was published …”, with the caveat that “An index published before 1931 in a work that is not consistently binominal is acceptable itself as a work consistent with the Principle of Binominal Nomenclature provided that the Principle is consistently applied to scientific names in the index; thus a scientific name published in such an index is available if the name satisfies the other provisions ….” (Article 11.4.3). As such, the scientific names in Seba’s (1759) work, even if they are interpreted as such, are not available as his work contravenes the said ICZN articles. Although Galil (1997) used these two Seba names as junior synonyms of C. calappa, on the basis of the current code, they also are also not available from her work even though she recognized them as available names. While the names Cancer flosculosus Seba, 1759, and Cancer heracleoticus Seba, 1759, are not available for nomenclatural purposes, it is nevertheless useful to determine what they are, especially since Seba’s plates are good. Another issue is that the Latin text in the index and the French tables that briefly describes the species and figures do not always match, often emphasising different aspects. As such, it is necessary to know what he wrote about the two species in both Latin and French. For Cancer flosculosus, the name was used for three different species crabs depicted in plate 19. The name was used in the contents page and subsequent index as follows: Unpaginated contents page for plate 19: Latin: “2. 3. Cancer flosculosus, pronus & supinus.” “13. Cancer flosculosus, Amboinensis, rarior, elegantissimus.” “24. Cancer flosculosus” French: 2. 3. “Cancre parfemé de fleurs, couché sur le dos & sur le ventre.” “13. Cancre d’Amboine, parfemé de felurs, très rare, d’une grande beauté.” “24. Cancre parfemé de fleurs.” Indices after plate 19: Latin “Num. 2. Cancer page 21 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan flosculosus, pronus. Tegmen superius, pallide rubens, coccineis maculis, tanquam flosculis, distinguitur Pedes antici, quorum sinister, unà cum forcipe süo, vel altero tanto major est, quam dexter, saturate rubri funt coloris. E capitis rotundi parte antica oculi protuberant. Reliqui decem pedes posteriores, pallide rubentes, prater quaternos, è quibus constant, articulos, longis quoque & acuminatis unguiculis gaudent. Num. 3. Idem supinus.” [Number 2. Cancer flosculosus, in prone position. The upper cover, pale red, with crimson spots, like flowers, is distinguished by chelipeds, whose left, together with its forceps, is much larger than the right, are of a saturated red color. Eyes protruded from the front of the round head. The remaining ten hind feet, pale red, besides the four, and of which they consist, the joints, also have long and pointed claws. Number 3. Same as above in supine position.] (Seba 1759: 46). French: “N°. 2. Cancre décoré de taches en façon de petites fleurs, vu par dessus. Sa coque du dessus du corps est d'un rouge-pâle, sursemé de tâches d'un rouge d'écarlate, qui ont quelque ressemblance à de petites fleurs. Ses deux bras sont d'un rouge-foncé; le gauche muni d'une bonne pince est de beaucoup plus grand que le droit. Ses deux yeux ronds sortens de la téte, où ils font places à la partie antérieure. Ses pieds font au nombre de dix, d'un rouge déteint, composés de quatre articulations, & munis d'ongles longs & pointus. N°. 3. Le méme Cancre couché sur le dos.” [Number 2. Crab decorated with spots in the shape of small flowers, seen from above. Its upper body shell is a pale red, dotted with scarletred spots, which have some resemblance to small flowers. Its two chelipeds are dark red; the left one fitted with good pincers is much larger than the right one. Its two round eyes protrude from the head, where they give way to the front part. Its feet are ten in number, a faded red, composed of four joints, and equipped with long and sharp claws. Number 3. The same crab lying on its back.] (Seba 1759: 46). Latin: “Num. 13: Cancer flosculosus, Amboinensis, rarior, elegantissimus. Postica ejus pars lata admodum utrinque acuminatos emittit processus, raro admodum in aliis Cancris conspicuos. Testa superior, dilutè Arantii coloris, ocellis exiguis, quos inter quidam minutissimi funt, tanquam squamulis vestita, parte antica saturate Arantio colore interstincta, singulari flosculorum quafi apparatu superbit. Oculi, anteriùs circa os proxime ad se mutuo collocati, ope articuli cujusdam prominent, ut eminus dispicere queant. Brachia crassa, forcipata, pedesque postici & cauda tam pressè sub testa superiore contracta hærent, ut vix videri possint. [Number 13: Cancer flosculosus, from Ambon, rare, very elegant. Its posterior part is very broad and, on both sides, there are pointed processes, very rarely visible in other Cancers. The upper shell, of a dilute orange color, with small eyes, some of which are the smallest, as if clothed with scales, the anterior part interspersed with a saturated orange color, boasts a unique arrangement of flower patterns. The eyes, placed close to each other near the front of the mouth, protrude by means of a joint, so that they can see from a distance. The chelipeds are thick, forked, and the hind feet and tail are so tightly contracted under the upper shell that they can scarcely be seen.] (Seba 1759: 47). French: “N°. 13. Cancre d' Amboine, rare, magnifique, sursemé de taches qui imitent des fleurs. Le dessus de son corps sort large sur le derriere, est encore remarquable par deux apophyses pointues qu'on trouve rarement dans d'autres Cancres. Sa coque ici orangée-clair, semble comme revêtue de petites écailles par les petits boutons dont elle est couvèrte, qu'on prendroit pour autant d'yeux, & parmi lesquels il y en a quantité d'une extrême petitesse; la partie antérieure de la coque est d'un orangé plus obscur, mais superbe par un assemblage de taches faites en façon de fleurs. Ses yeux font situés l'un près de l'autre de chaque côté de la bouche, s’avançant en avant par le moyen de quelque appendice pour voir de loin. Ses deux gros bras sourchus, ses jambes, & sa queuë font si sort ramassés sous la coque, qu'on peut à peine les découvrir.” [Number 13. Ambon Crab, rare, magnificent, dotted with spots that imitate flowers. The top of its body comes out wide at the back, is further remarkable by two pointed processes which are rarely found in other Dunces. Its shell, here light orange, seems to be covered in small scales by the small buttons with which it is covered, which we take for so many eyes, and among which there are a number of extremely small ones; the anterior part of the shell is a darker orange, but superb with an assembly of spots made in the shape of flowers. Its eyes are located close to each other on each side of the mouth, projecting forward by means of some appendage to see from a distance. Its two big chelipeds, its legs, and its pleon are so compacted page 22 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan under the hull that we can barely discover them.] (Seba 1759: 47). Latin: “Num. 24. Cancer flosculosus, pronus. Pallida hîc rubedo saturatius rubris quasi florum foliolis, hinc inde dispersis, variegatur. Postica pars latio est, antica angustior rotunde convergit. Inter oculos apice quodam os prominet. Brachia crassa sunt; pedes vero postici minores sub ventre delitescunt.” [Number 24. Cancer flosculosus, in prone position. Carapace saturated with pale red spots reminiscent of flowers blooming. The posterior part is broad, the front is narrower and roundly converging. Between the eyes a kind of mouth protrudes at the tip. The chelipeds are thick; but the smaller hind feet are hidden under the belly.] (Seba 1759: 49). French: “N°. 24. Cancre orné de taches faites en façon defleurs, vu par dessus. Il est d’un rouge incarnat. émaillé de taches d'un rouge plus foncé, faites en maniere de petites feuilles de fleur femées çà & là. Sa partie de derriere est large, & sa partie de devant plus étroite & plus arrondie. Sa bouche pointue s'avance entre les yeux. Ses bras font gros. Ses pieds font petits, cachés fous le ventre.” [Number 24. Crab decorated with spots made in the style of flowers, seen from above. It is a crimson red. enamelled with spots of a darker red, made in the manner of small flower leaves cut here and there. Its back part is wide, and its front part narrower and more rounded. His pointed mouth juts out between his eyes. His chelipeds are big. His feet are small, hidden beneath his ventrum.] (Seba 1759: 49). For Cancer heracleoticus, the name was used for a crab depicted in plate 20. The name was used in the contents page and subsequent index as follows: Unpaginated contents page for plate 20: Latin: “7. 8. Cancer Heracleoticus, pronus and supinus.” French: “7. 8. Cancre héracléotique, vu par dessus & par dessous.” Indices after plate 20: Latin: “Num. 7. Cancer Heracleoticus, pronus. Rarissimus hic Cancer, unà cum aliis speciminibus, ex Amboina mihi missus est, sub nomine Cancri cristati, seu Galli marini. Aristoteli Pagurus vocatur ex mari Adriatico; Bellonio Cancer Heracleoticus, species Majæ; J. Cæsari Scaligero Pagurus, crista galli; Rondeletio Pagurus Migrane, id est, Malum Punicum; & Rumphio, in Museo Amboinensi, Cancer Calappoides. Unde equidem pater, aliis Authoribus alia placuisse nomina, pro varia similitudine, quam invenisse sibi quisque imaginatus est. Parte postica latus est limbo elato instructus. Cauda brevis & obrusa: amica pars angustior rotundè convergit. Testa superior, antrorsum, malum Punicum, maturescens, ipso etiam colore refert. Posteriora versus latior testa, instar aquæ lente fluctuantis sulcata, undas maris dense congestas repræsentat. Oculi profunde intra testaceas orbitas, ad latera nasi veluti essictas, delitescunt. Num. 8. Idem supinus. Brachia duo antica, Galli capitibus fimilia, anterius admodum lata, incurvum singula unguem gerunt acuminatum, rubicundum, qui cum inferna forcipis parte, itidem antrorsum late protensa, arctè committitur limbo interno dentatus. Testa superior, geminæ instar galli cristæ, serrata & quam elegantissime picta est. Pedes octo postici, dilute flavi, ungues acutos, rubentes, gerunt.” [Number 7. Cancer Heracleoticus, in prone position. This very rare Cancer, together with other specimens, was sent to me from Ambon, under the name of Crested Cancer, or Rooster of the Sea. Aristotle’s Pagurus is called from the Adriatic sea; Bellonio’s Cancer Heracleoticus, a species of Majæ; J. Caesar Scaligero’s Pagurus with a cock’s crest; Rondeletio’s Pagurus Migrane, that is, Phoenician Apple [pomegranate]; & Rumphius, in the Museum of Ambon, Cancer Calappoides. Wherefore, indeed, the originator, each one imagined that other authors had chosen other names, for the various likenesses which he had found for himself. The rear side is equipped with a raised rim. The tail is short and stubby: the friendly part is narrower and rounded. The upper shell, in front, represents the Punic apple, ripening, in the same color. The posterior towards the wider shell, like the furrows of slowly undulating water, represents the densely packed waves of the sea. The eyes are hidden deep within the orbits, along the sides of the front. Number 8. The same in supine position. Two chelipeds in front, similar to rooster heads, very wide in front, each curved claw is pointed, red, forming very sharp forceps on the side, also extended wide along the front is a serrated inner rim. The upper shell is like a cock's crest, serrated and most elegantly coloured. The hind feet are eight, pale yellow, and have sharp, red claws.] (Seba 1759: 51–52). French: “N°. 7. Cancre Héracléotique, vu par dessus. Ce Cancre qui est extrêmement rare m'a été envoyé d'Amboine avec d’autres Curiosités, sous le nom de Cancre portant une Crête, ou page 23 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Coq de Mer. Aristote l'appelle le Squinado de la Mer Adriatique, Belon le Cancre Héracléotique Espece de Mæas, J. César Scaliger le Squinado à crête de Coq, Rondelet le Cancre Migraine, c'està-dire ressemblant à une Grenade qu'on nomme Migraine en Languedoc. Ainsi chaque Auteur lui a donné un nom à fa fantaisie, conformément à la ressemblance qu'il lui a trouvée avec telle ou telle chose. Il est large par derriere, avec une bordure relevée. Sa queuë est courte & obtuse. La partie antérieure de son corps est étroite & arrondie. La coque du dessus du corps ressemble par devant de couleur & de figure à une Grenade presque mure; ensuite s'élargissant sur le derriere du corps, elle est fillonnée de rayes faitesen ondes, representant celles de l'eau quand elles se touchent de prés & se succedent lentement les unes aux autres. Ses yeux sont cachés dans leurs orbites, qui semblent être creusés vers les côtés du nez. No. 8. Le même Cancre, vu par dessous. Ses deux bras ressemblent par leur figure à des tétes de Coq. Ils sont fort larges dans la partie antérieure, armés de pinces pointues au bout, de couleur rouge, dentelées intérieurement, & dont la supérieure qui est crochue entre dans l'inférieure par engrénure. La coque qui couvre l'Animal est ornée de belles couleurs, & entaillée sur les bords d'une dentelure qui imite une double créte de Coq.” [Number 7. Cancer Heracleoticus, seen from above. This crab which is extremely rare was sent to me from Ambon with other Curiosities, under the name of Crab wearing a Crest, or Sea Cock. Aristotle calls it the Squinado of the Adriatic Sea, Belon the Heracleotic Crab Species of Mæas, J. César Scaliger the Cock-crested Squinado, Rondelet the Cancer Migraine, resembling a Pomegranate which is called Migraine in Languedoc. Thus each Author has given it a name at will, in accordance with the resemblance he found to be close with this or that thing. It is wide from behind, with a raised border. Its tail is short & obtuse. The front part of its body is narrow & rounded. The shell of the upper body resembles an almost ripe pomegranate from the front in colour and figure; then widening on the back of the body, it is filled with wave-like stripes, like those of water when they touch closely and slowly follow one another. Its eyes are hidden in their sockets, which appear to be sunken towards the sides of the nose. Number 8. The same crab, seen from below. Its two arms look like rooster heads. They are very wide in the front part, armed with pointed pincers at the end, red in color, serrated internally, and the upper one, which is hooked, enters the lower one. The shell which covers the animal is decorated with beautiful colors and notched on the edges with a serration which imitates a double crest of a rooster.] (Seba 1759: 51–52). Seba’s (1759) “Cancer flosculosus” is a mix of three species. The specimen depicted in plate 19 figures 2 and 3 (no locality given) is clearly what is today called Carpilius convexus (Forskål, 1775) (Carpiliidae), a widespread species in the Indo-West Pacific. That in plate 19 figure 13 is clearly Calappa flammea (Herbst, 1794), a common species in the Atlantic. Although Seba reports that the specimen was from Ambon, there must be a mistake in the labelling. As for the specimen shown in plate 19 figure 24 (no location), it is probably a young Calappa calappa as its carapace has low nodules on the carapace. Seba’s (1759) “Cancer heracleoticus” (no location indicated) is clearly C. calappa, the morphology and colour agreeing in all aspects. Calappa capellonis Laurie, 1906 # (Figs. 6, 35A) Calappa capellonis — present record. Taxonomy: Takeda and Koyama (1974), Chen (1993), Galil (1997), Ng (2002a). Type: Syntypes NHM 1907.5.22.13-15 (Galil 1997). Type locality: Gulf of Mannar, Sri Lanka. Distribution: Western Indian Ocean to West Pacific: from Madagascar, Seychelles, to Indonesia, South China, Taiwan and Japan, to Australia and New Guinea (Galil 1997). Habitat: Subtidal; rocky substrates of coarse sand or rock fragments; depths 9 to 80 m (Galil 1997). Remarks: The present species was described as a variety of Calappa gallus (Herbst, 1803), but is distinguished from it by more flattened and smoother nodules on the carapace among various characters (Laurie 1906). The distinction was dismissed by Rathbun (1937), but later recognized by Takeda and Koyama (1974) who raised it to a full species (see also Chen 1993; Galil 1997). This species is a new record for the fauna of Hong Kong. Calappa hepatica (Linnaeus, 1758) # Calappa calappa — Hill and Phillipps 1981: 156, pl. 52k [not Cancer calappa Linnaeus, 1758]. Calappa hepatica — Morton and Morton 1983: 157, fig. 10.4(4). Taxonomy: Chen (1993), Galil (1997), Chen and Sun (2002). page 24 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Type: Status unknown. Type locality: “in Indiis”: Indian Ocean. Distribution: Western Indian Ocean to East Pacific: South Africa, Madagascar, Red Sea, Persian/ Arabian Gulf, to South China Sea and East Asia, Indonesia, Papua New Guinea, Australia, New Zealand and New Caledonia, to Hawaii and Clipperton Island (Galil 1997); exotic species in the Mediterranean (Brockerhoff and McLay 2011; Galil 2011). Habitat: Intertidal to subtidal; substrates sandy, muddy or gravelly, to depths of 100 m (Galil 1997). Calappa lophos (Herbst, 1782) # Calappa lophos — Wong et al. 2021: 8, fig. 5, pl. 1D. Taxonomy: Chen (1993), Ng et al. (1999a), JCY Lai et al. (2006). Type: Lectotype ♂ ZMB 2168 (K Sakai 1999; JCY Lai et al. 2006). Type locality: East Indian Seas. Distribution: Western Indian Ocean to West Pacific: from South Africa, Andaman Sea, South and East China Seas, Taiwan, Japan, and Australia (Chen and Sun 2002; Davie 2002; JCY Lai et al. 2006). Habitat: Subtidal; substrates of muddy sand, fine sand or shell fragments; depths 30 to 100 m (Chen and Sun 2002). Calappa philargius (Linnaeus, 1758) # Calappa cristata — Stimpson 1858c: 162; Stimpson 1907: 165. Calappa philargius — Gee 1926: 160; Shen 1931a: 104, text-figs. 10, 11, pl. 8; Shen 1940a: 214; Hill et al. 1978: 114; Morton 1979b: 136, tab. 7.1; Hill and Phillipps 1981: 156, pl. 52j; Morton and Morton 1983: fig. 10.4(6); Davie 1992b: tab. 1; Galil 1997: 307, figs. 17d, 20d, 21, 23; Morton and Ruxton 1997: 1 unnumb. fig. (51); Blackmore and Rainbow 2000: app. 1; B Chan and Leung 2002: 25, fig. 2G; Leung et al. 2004: 87, 1 unnumb. fig.; Tsang et al. 2020: 68, 2 unnumb. figs.; Wong et al. 2021: 8, fig. 6, pl. 1E. Taxonomy: Chen (1993), Galil (1997), Chen and Sun (2002), Ng et al. (2002). Type: Neotype ♂ ZRC 1998.67 (designated by Ng et al. 2002). Type locality: Changi Beach, Singapore (neotype). Distribution: Eastern Indian Ocean to West Pacific: from Straits of Malacca eastwards, to Southeast and East Asia, and Western Australia (Ng et al. 2002). Habitat: Subtidal; substrates sandy-muddy, shelly, of depth 24 to 100 m (Chen and Sun 2002). Fig. 6. Calappa capellonis Laurie, 1906 (female, 62.2 × 47.4 mm, off Kau Sai Chau): A, carapace; B, right chela; C, left chela. page 25 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Habitat: Freshwater; low-gradient streams with muddy substratum (Ng and Dudgeon 1992); occasionally seen in freshwater marshes. Remarks: Records under the name Parathelphusa sinensis H. Milne Edwards, 1853, and even the type series (from “Mers de la Chine”), was reported to be heterogeneous (Ng and Dudgeon 1992). A lectotype for P. sinensis was designated by Ng and Dudgeon (1992), and another distinct species in the former type series of P. sinensis was described as Somanniathelphusa falx. Old material from Hong Kong were referred to the present species in the same paper. Superfamily Goneplacoidea Macleay, 1838 Family Euryplacidae Stimpson, 1871 Eucrate alcocki Serène, in Serène & Lohavanijaya, 1973 # ? Eucrate crenata var. dentata — Alcock 1900b: 301 [not Heteroplax dentatus Stimpson, 1858]. Eucrate alcocki — Shin and Thompson 1982: app.; Morton and Morton 1983: 98; Shin 1989: app. 1; Ong Che and Morton 1991: tab. 1; Blackmore and Rainbow 2000: app. 1; SY Lee et al. 2000: tab. 3; Castro and Ng 2010: 18, figs. 2C, 4A–C, 14A–C; Wong et al. 2021: 12, fig. 15, pl. 3C. Taxonomy: Serène and Lohavanijaya (1973), Castro and Ng (2010). Type: Holotype ♂ MNHN-B10134 and paratype ♀ in MNHN (Castro and Ng 2010). Type locality: Nhatrang Bay, Vietnam. Distribution: South China Sea and East Asia: Singapore, eastern Peninsular Malaysia, Gulf of Thailand, Vietnam, the Philippines and South and East China Seas (Castro and Ng 2010). Habitat: Subtidal; substrates muddy-sandy; to depths of 50 m (Dai and Yang 1991). Eucrate crenata (De Haan, 1835) # Pilumnoplax sulcatifrons Stimpson, 1858b: 93; Stimpson 1907: 90. Eucrate crenata — Alcock 1900b: 300; Gordon 1931: 528; Shen 1940a: 229; Shin and Thompson 1982: app.; Thompson and Horikoshi 1982: tab. 1; Thompson et al. 1982: tab. 1; Thompson and Shin 1983: tab. 4; Shin 1990: tab. 2; Ong Che and Morton 1991: tab. 1; Davie 1992b: tab. 1; SY Lee and Leung 1999: 78; Blackmore and Rainbow 2000: app. 1; SY Lee et al. 2000: tab. 3; Castro and Ng 2010: 21, figs. 2A, B, 3A–G, 14D–F; Wong et al. 2021: 12, fig. 16, pl. 3D, E. Heteroplax sulcatifrons — Gee 1926: 163 [list]. Eucrate sulcatifrons — Gordon 1931: 528; Shen 1940a: 230; Davie 1992b: tab. 1. Taxonomy: Castro and Ng (2010). Type: Lectotype ♂ RMNH D 287 and paralectotypes in RMNH (designated by Yamaguchi and Baba (1993); see also Fransen et al. (1997)). Type locality: Japan, locality unspecified. Distribution: Western Indian Ocean to South China Sea and East Asia: questionably Madagascar, Seychelles and northern Australia, verified from the Red Sea, Persian/Arabian Gulf, Arabian Sea, India, Southeast and East Asia (Castro and Ng 2010). Habitat: Subtidal; to depths of 36 m (Castro and Ng 2010). Eucrate solaris Yang & Sun, 1979 Eucrate solaris — SY Lee et al. 2000: tab. 3. Taxonomy: Chen (1998), Hsueh and Huang (2002), Castro and Ng (2010). Type: Holotype ♂ BNHM J79136. Type locality: Beihai, Guangxi, China. Distribution: West Pacific: Gulf of Thailand, the Philippines, South China Sea, Taiwan, Vanuatu; questionably from Ryukyu Islands, Japan (Castro and Ng 2010). Habitat: Subtidal; of muddy-sandy bottom (Dai and Yang 1991); to depths of at least 200 m but with records from tangle nets reaching to 500 m (Castro and Ng 2010). Eucrate tripunctata Campbell, 1969 # (Figs. 8, 35C–E) Eucrate tripunctata — present record. Taxonomy: Campbell (1969), Castro and Ng (2010). Type: Holotype ♂ QM W3034 and paratypes in QM. Type locality: Moreton Bay, Queensland, Australia. Distribution: Eastern Indian Ocean to West Pacific: from Queensland Australia), Mergui Archipelago, Singapore, Gulf of Thailand (Castro and Ng 2010) to South China (Hong Kong). Habitat: Intertidal to shallow subtidal; substrates of coarse sand and shell and rock fragments. Remarks: In the recent revision of the family, Castro and Ng (2010) included a live-color photograph under presentation of E. crenata, entirely pale orange in color, from intertidal habitats of Hong Kong (fig. 2B; Pl. 1C). From thousands of trawled specimens our colleagues has sorted, none display such coloration. Based on examined material from local shores, given the markedly broad P5 propodus (Fig. 8B), as well as obtuse anterolateral teeth (1 to 3; Fig. 8A) (cf. Campbell 1969; Castro and Ng 2010), we have identified them as page 32 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan E. tripunctata instead. This species is a new record for the fauna of Hong Kong. Henicoplax nitida (Miers, 1879) Henicoplax nitida — Castro and Ng 2010: 63, figs. 23A–D, 24G, H. Taxonomy: Castro and Ng (2010). Type: Holotype ♀ NHM 78.11 (Castro and Ng 2010). Type locality: Strait of Korea. Distribution: East Asia: South China (Hong Kong) and Japan. Habitat: Shallow subtidal (Castro and Ng 2010). Heteroplax transversa Stimpson, 1858 # Heteroplax transversus Stimpson, 1858b: 94; Stimpson 1907: 95. Heteroplax transversa — Gee 1926: 163 [list]; Castro and Ng 2010: 73, figs. 27G–I, 28A–F; Wong et al. 2021: 12, fig. 17, pl. 3F. Eucrate transversa — Gordon 1931: 528; Shen 1940a: 229. Heteroplax nagasakiensis — Ong Che and Morton 1991: tab. 1. Taxonomy: Castro and Ng (2010). Type: Presumably lost. Type locality: Hong Kong. Distribution: East Asia: South China and Japan (Castro and Ng 2010). Habitat: Subtidal; to depths of 18 to 50 m (Castro and Ng 2010). Trissoplax dentata (Stimpson, 1858) # Heteroplax dentatus Stimpson, 1858b: 94; Stimpson 1907: 94. Heteroplax dentata — Gee 1926: 163 [list; “near Hong Kong”]. Eucrate dentata — Gordon 1931: 528; Shen 1940a: 229; Davie 1992b: tab. 1. Eucrate costata — Blackmore and Rainbow 2000: app. 1. Trissoplax dentata — Castro and Ng 2010: 98, figs. 39D–F, 40A–F, 41A–G, 43A–C; Wong et al. 2021: 13, fig. 18, pl. 4A. Taxonomy: Castro and Ng (2010). Type: Neotype ♂ QM W27400 (designated by Castro and Ng 2010). Type locality: Tolo Harbour, New Territories, Hong Kong (neotype). Distribution: Western Indian Ocean to West Pacific: from South Africa, Persian/Arabian Gulf, India, Singapore, Thailand to South China, Indonesia and Australia (Castro and Ng 2010). Habitat: Shallow subtidal to depths of 40 m (Castro and Ng 2010). Fig. 8. Eucrate tripunctata Campbell, 1969 (male, 39.6 × 31.0 mm, Lung Mei): A, carapace; B, right P5; C, pleon. page 33 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Family Goneplacidae Macleay, 1838 Subfamily Goneplacinae Macleay, 1838 Carcinoplax haswelli (Miers, 1884) # Carcinoplax sinica — Wong et al. 2021: 14, fig. 20, pl. 4C. Taxonomy: Chen (1984: as C. sinica), Ng and Mitra (2019: as C. sinica), Ng et al. (2022a). Type: Lectotype ♂ and paralectotype ♀ in NHM 1882.7 (designated by Ng et al. 2022a). Type locality: Arafura Sea, Australia. Distribution: West Pacific: Southern Taiwan and South China Sea, to northern Australia and Papua New Guinea (Ng et al. 2022a). Habitat: Subtidal; substrates sandy and stony (Dai and Yang 1991); depths 25 to 187 m, usually less than 100 m (Ng et al. 2022a). Remarks: The better understood C. sinica Chen, 1984 has been recorded from various localities in vicinity of the South China Sea (Castro 2007; Ng and Mitra 2019). Ng et al. (2022a) recently demonstrated that a poorly known Homoioplax haswelli (Miers, 1884) from northern Australia, the syntypes being juveniles, was a senior synonym of C. sinica. The genetics of this and allied species was recently done by Prema et al. (2024). Carcinoplax longimana (De Haan, 1833) Carcinoplax longimanus — Gordon 1931: 528; Shen 1940a: 229. Taxonomy: Chen (1984), Guinot (1989), Castro (2007). Type: Lectotype ♂ RMHN D 308 and paralectotypes in RMNH (designated by Yamaguchi and Baba 1993; see Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: Western Indian Ocean to South China Sea and East Asia: from South Africa, Madagascar, to South and East China Seas and Indonesia (Castro 2007). Habitat: Subtidal; from depths 66 to 377 m (Castro 2007). Carcinoplax purpurea Rathbun, 1914 # Carcinoplax purpurea — Davie 1992b: tab. 1; Wong et al. 2021: 14, fig. 19, pl. 4B. Taxonomy: Chen (1984), Guinot (1989), Castro (2007). Type: Holotype ♀ USNM 46143. Type locality: Between Luzon and Mindoro, Verde Island Passage, the Philippines. Distribution: Southeast and East Asia: from Indonesia to the Philippines, Vietnam, South and East China, Taiwan and Japan (Castro 2007). Habitat: Subtidal; depths from 17 to 180 m (Castro 2007). Entricoplax vestita (De Haan, 1833) Carcinoplax vestita — Blackmore and Rainbow 2000: app. 1. Taxonomy: Chen (1984: as Carcinoplax), Castro (2007). Type: Lectotype ♀ RMHN D 311; paralectotypes in RMHN and MNHN (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: East Asia: East China Sea and Yellow Sea, to Pacific coast of Japan (Castro 2007). Habitat: Subtidal; substrates muddy and sandy (Dai and Yang 1991); depths 15 to 110 m (Castro 2007). Family Scalopidiidae Števčić, 2005 Scalopidia spinosipes Stimpson, 1858 # Scalopidia spinosipes Stimpson, 1858b: 95; Alcock 1900b: 325 [part: material from Hong Kong only]; Gordon 1931: 528; Rathbun 1931: 100 [“near Hong Kong”]; Shen 1940a: 230; AC Evans 1967: 407; Davie 1992b: tab. 1; SY Lee et al. 2000: tab. 3; Ng and Castro 2013: 60, figs. 1, 2A–D, 3, 5A, 6A, 7A–D, 8A, B, 9A, B, 10A, B, 11A, B, 12A–D, 14A; Wong et al. 2021: 143, fig. 21, pl. 4D. Hypophthalmus leucochirus Richters, in Lenz & Richters, 1881: 429, pls. 1–10. Taxonomy: Ng and Castro (2013). Type: Lectotype ♂ and paralectotype ♀ in NHM (Stimpson Collection NHM 61.44) (designated by Ng and Castro 2013). Type locality: Hong Kong. Distribution: Southeast and East Asia: Singapore, Indonesia, the Philippines, Gulf of Thailand, to Vietnam and Hong Kong (Ng and Castro 2013). Habitat: Subtidal; substrates muddy-sandy; depths about 20 m (Dai and Yang 1991). Superfamily Hexapodoidea Miers, 1886 Family Hexapodidae Miers, 1886 Hexalaughlia orientalis (Rathbun, 1909) # Hexalaughlia orientalis — Ng and Wong 2019: fig. 1; Wong et al. 2021: 14, fig. 22, pl. 4E. Taxonomy: Rathbun (1910), Guinot (2006). Type: Probably in now NHMD (original text), yet page 34 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan “not possible to examine” as noted by Guinot (2006). Type locality: North of Koh Kut, eastern Gulf of Thailand. Distribution: South China Sea and East Asia: Gulf of Thailand, South China, Taiwan, and possibly also to Japan (Guinot 2006; Manning and Holthuis 1981). Habitat: Subtidal; substrates muddy, commensal in tubes of annelids (Dai and Yang 1991). Hexapinus simplex Rahayu & Ng, 2014 # Hexapinus simplex — Ng and Wong 2019: 236, fig. 2; Wong et al. 2021: 14, fig. 23, pl. 4F. Taxonomy: Rahayu and Ng (2014), Naruse et al. (2017). Type: Holotype ♂ MZB-Cru 3790; paratypes in MZB, USNM and ZRC. Type locality: Ekas, Lombok, Indonesia. Distribution: South China Sea and East Asia: Indonesia, Singapore, to South China and Taiwan Strait, and Ryukyus (Japan) (Rahayu and Ng 2014). Habitat: Intertidal and subtidal; from muddy or sandy substrates, or intertidal seagrass beds; has also been reported from tubes of chaetopterid worms (Rahayu and Ng 2014). Mariaplax chenae Rahayu & Ng, 2014 # ? Hexapus anfractus — Morton 1988: 36, pl. 7 [not Lambdophallus anfractus Rathbun, 1909]. ? Hexapus granuliferus — Ong Che and Morton 1991: tab. 1 [not H. granuliferus Campbell & Stephenson, 1970]. Mariaplax chenae — Ng and Wong 2019: 237, figs. 3–5; Wong et al. 2021: 15, fig. 24, pl. 5A. Taxonomy: Rahayu and Ng (2014), Naruse et al. (2017), Ng and Wong (2019). Type: Holotype ♂ in IOCAS; paratypes in IOCAS and ZRC. Type locality: Coastal waters of Zhejiang, China. Distribution: East Asia: coasts of China to Shandong, and Japan (Rahayu and Ng 2014). Habitat: Subtidal; from depths of 4 to 39 m (Rahayu and Ng 2014). Mariaplax exigua Ng & Wong, 2019 # Mariaplax exigua Ng & Wong, 2019: 239, figs. 6, 7; Wong et al. 2021: 16, fig. 25, pl. 5B. Taxonomy: Ng and Wong (2019). Type: Holotype ♂ ZRC 2018.0706. Type locality: Tolo Channel, Hong Kong. Distribution: So far from Hong Kong only. Habitat: Subtidal; shallow waters. Superfamily Hymenosomatoidea Macleay, 1838 Family Hymenosomatidae Macleay, 1838 Subfamily Hymenosomatinae Macleay, 1838 Halicarcinus setirostris (Stimpson, 1858) Rhynchoplax setirostris Stimpson, 1858b: 109; Stimpson 1907: 148; Shen 1940a: 231 [no new specimens]. Taxonomy: Shen (1932b: as H. yangi), Sakai (1938: as Rhynchoplax), Ng et al. (1999b). Type: Presumably lost. Type locality: Hong Kong. Distribution: East Asia: coasts of China (South China Sea to Yellow Sea), Taiwan and Japan (Ng et al. 1999b; Ng and Jeng 1999; Yang et al. 2008). Habitat: Subtidal; substrate muddy or shelly; depths 50 to 100 m (Dai and Yang 1991). Neorhynchoplax introversa (Kemp, 1917) # (Figs. 9, 35F) Neorhynchoplax introversa — present record. Taxonomy: Kemp (1917: as Rhynchoplax), Ng et al. (1999b). Type: Holotype ZSI 9730/10 (original text). Type locality: Tai Hu Lake, Jiangsu (as Kiangsu), China. Distribution: Coasts of China: from estuaries of Pearl River to Yellow Sea. Habitat: Intertidal to freshwater; under rocky fragments along intertidal mudflats, or among foliage on freshwater wetlands. Remarks: This species is a new record for the fauna of Hong Kong. Neorhynchoplax sinensis (Shen, 1932) # (Figs. 10, 36A) Neorhynchoplax sinensis — present record. Taxonomy: Shen (1932b: as Rhynchoplax), Ng et al. (1999b), Takeda (2001: as Elamenopsis). Type: Holotype ♂ ZMFMIB 8615: status unknown. Type locality: Lutao, Shandong, China. Distribution: Coasts of China: from Hainan to Yellow Sea. Habitat: Intertidal; under rock fragments, along crevices, or among oyster clumps. Remarks: This species is a new record for the fauna of Hong Kong. page 35 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Superfamily Leucosioidea Samouelle, 1819 Family Iphiculidae Alcock, 1896 Iphiculus spongiosus Adams & White, 1849 # Iphiculus spongiosus — Stimpson 1858c: 161; Stimpson 1907: 159, pl. 18(8); Gee 1926: 160 [list; “near Hong Kong”]; Shen 1940a: 215; Ong Che and Morton 1991: tab. 1; Davie 1992b: tab. 1; Wong et al. 2021: 16, fig. 26, pl. 5C. Taxonomy: Serène (1955), Chen (1989), Chen and Sun (2002). Type: Syntypes probably in NHM (NHM Data Portal). Type locality: The Philippines, locality unspecified. Distribution: Western Indian Ocean to South China Sea and East Asia: from Red Sea, India, Singapore, Indonesia, South and East China Seas and Japan (Chen 1989). Habitat: Subtidal; bottom of sandy mud of soft mud, depths of 11 to 204 m (Chen 1989). Pariphiculus mariannae (Herklots, 1852) Pariphiculus mariannae — Gordon 1931: 527; Shen 1940a: 215; Guinot 1979: 147, pl. 15(8–10); CGS Tan 1996: 1049 [21°44'N, 114°48'E]. Taxonomy: Serène and Lohavanijaya (1973), Chen (1989), Chen and Sun (2002), Y-J Shih et al. (2017). Type: Syntypes in “Musée de Louvain” (original text); whereabouts and status unknown (see Fransen et al. 1997: 212 on the museum). Type locality: China, locality unspecified. Distribution: Western Indian Ocean to South China Sea and East Asia: Red Sea, Bay of Bengal, the Philippines, Indonesia, and South China Sea to Japan (Chen 1989). Habitat: Subtidal; substrates of mud, sandy mud; depths 65 to 296 m (Chen 1989). Fig. 9. Neorhynchoplax introversa (Kemp, 1917) (male, 3.2 × 3.4 mm, Mai Po Marshes): A, carapace; B, frontal region and maxilliped 3, ventral view; C, cheliped carpus and chela; D, right P5; E, pleon; F, right G1, ventral view; G, tip of right G1, ventral view. Structures A to D denuded. page 36 of 155 Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Family Leucosiidae Samouelle, 1819 Subfamily Cryptocneminae Stimpson, 1907 Onychomorpha lamelligera Stimpson, 1858 Onychomorpha lamelligera Stimpson, 1858c: 162; Stimpson 1907: 164, pl. 19(8, 8a); Gee 1926: 161 [list]; Shen 1940a: 217 [no new specimens]. Taxonomy: Serène and Soh (1976), Chen and Sun (2002). Type: Presumably lost. Type locality: Hong Kong. Distribution: Eastern Indian Ocean to South China Sea: Palk Strait, Phuket (Thailand), Singapore, Gulf of Thailand, the Philippines and South China (Chen and Sun 2002). Habitat: Subtidal; substrates soft mud, sandy mud or sandy shelly, depths 6 to 27 m (Chen and Sun 2002). Subfamily Ebaliinae Stimpson, 1871 Alox somphos C.G.S. Tan & Ng, 1995 # (Figs. 11, 36B) Alox somphos — present record. Taxonomy: Serène (1954: as Oreophorus (O.) rugosus), CGS Tan and Ng (1995), Galil and Ng (2007). Type: Holotype ♀ ZRC-1984.5644, and paratypes in ZRC and ZMA. Type locality: Sentosa Reef, Singapore. Distribution: Southeast and East Asia: Indonesia, Palau, Singapore, Vietnam (CGS Tan and Ng 1995), and now South China (Hong Kong). Habitat: Lower intertidal and subtidal; under rock fragments, in rocky or coral reefs. Remarks: This interesting species was identified by a slender, gently tapering G1 (Fig. 11E, F) as Fig. 10. Neorhynchoplax sinensis (Shen, 1932) (male, 2.7 × 3.0 mm, Lower Pak Nai): A, carapace; B, maxilliped 3; C, left P5; D, pleon; E, right G1, mesial view; F, tip of right G1, mesial view. Structures A to D denuded. page 37 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan illustrated by Galil and Ng (2007: fig. 5C, D), as well as overall form of carapace (Figs. 11A, 38B; CGS Tan and Ng 1995: pl. 8D). This is a new record for the fauna of Hong Kong. Arcania globata Stimpson, 1858 Arcania globata — Stimpson, 1858c: 160; Stimpson 1907: 156, pl. 18(9); Gee 1926: 160 [list]; Shen 1940a: 215. Taxonomy: Shen (1937a), Galil (2001a), Chen and Sun (2002). Type: Presumably lost. Type locality: China Seas at 23°N (near Hong Kong). Distribution: East Asia: South China Sea to Yellow Sea, Korea and Japan (Galil 2001a; Chen and Sun 2002). Habitat: Subtidal; substrates sandy, muddy or shelly; depths 30 to 150 m (Chen and Sun 2002). Arcania gracilis Henderson, 1893 Arcania quinquespinosa — Gordon 1931: 527; Shen 1940a: 215; CGS Tan 1996: 1028, figs. 2b, e, 7F [21°42–46'N, 114°47–50'E]. Arcania gracilis — Galil 2001a: 184, figs. 2D, 5D. Taxonomy: Chen (1989: as A. quinquespinosa), CGS Tan (1996: as A. quinquespinosa), Galil (2001a), Chen and Sun (2002: as A. quinquespinosa). Type: Syntype ♂ NHM 1884.34 (Galil 2001a). Type locality: Gulf of Martaban, Myanmar. Distribution: Western Indian Ocean to West Pacific: from Madagascar, Red Sea, Persian/Arabian Gulf, India, to Southeast and East Asia, Australia, New Caledonia and Vanuatu (Galil 2001a). Habitat: Subtidal; substrates of sand, mud, shell fragments and coarse sand; depths 21 to 185 m (Chen and Sun 2002). Fig. 11. Alox somphos C.G.S. Tan & Ng, 1995 (male, 14.7 × 9.9 mm, Shelter Island): A, carapace; B, maxilliped 3; C, chela, right; D, thoracic sternum, showing left G1; E: pleon; F: left G1, mesial view; tip of left G1, mesial view. page 38 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Arcania heptacantha De Man, 1907 # Iphis septemspinosa — Stimpson 1858c: 161; Stimpson 1907: 157 [not Cancer septemspinosa Fabricius, 1787]. Arcania heptacantha — Hill 1982: 201, pl. 4C; Shin and Thompson 1982: app.; Davie 1992b: tab. 1; Blackmore and Rainbow 2000: app. 1; SY Lee et al. 2000: tab. 3; Galil 2001a: 187, figs. 2D, 6A; Wong et al. 2021: 16, fig. 27a–c, pl. 5D. Taxonomy: De Man (1907), Galil (2001a), Chen and Sun (2002). Type: Holotype ♀ RMNH D 42112 (Fransen et al. 1997). Type locality: Unknown (Herklots 1861 (Iphis heptacantha as nomen nudum); Galil 2001a). Distribution: South China Sea and East Asia: Singapore, South and East China Seas, Taiwan and Japan (Galil 2001a). Habitat: Subtidal; substrate soft-muddy or muddysandy; depths 6 to 150 m (Chen and Sun 2002). Arcania septemspinosa (Fabricius, 1787) # Arcania septemspinosa — Miers 1886: 300; Koelbel 1897: 710; Balss 1922a: 132; Gee 1926: 160 [list; “near Hong Kong”]; Gordon 1931: 527; Shen 1940a: 215; CGS Tan 1996: 1029, figs. 1D, 2a [21°42'N, 114°50'E]; Galil 2001a: 192, figs. 3B, 7B; Wong et al. 2021: 16, fig. 27d, e, pl. 5E. Taxonomy: Chen (1989), Galil (2001a), Chen and Sun (2002). Type: Syntypes in ZMK (Zimsen 1964), in present NHMD (examined by second author, not studied by Galil 2001a). Type locality: Indian Ocean. Distribution: Western Indian Ocean to Central Pacific: from South Africa, Madagascar, Red Sea, Persian/Arabian Gulf, India, to Southeast and East Asia, and northern Australia and Fiji (Galil 2001a). Habitat: Subtidal; substrate muddy sand or soft, muddy; depths 21 to 180 m (Chen and Sun 2002). Arcania tropicalis Naruse, 2014 Arcania tropicalis Naruse, 2014: 320, figs. 18–21. Taxonomy: Naruse (2014). Type: Holotype ♂ NTOU B00111, and paratypes in NTOU, ZRC, NMMB. Type locality: Kezailiao, Kaohsiung, Taiwan. Distribution: West Pacific: Vanuatu, the Philippines, South China (Hong Kong), and Taiwan. Habitat: Subtidal; collected by trawling or tangle nets; to depths of 500 m (Naruse 2014). Arcania undecimspinosa De Haan, 1841 # Arcania undecimspinosa — Shen 1931a: 107, pl. 10(1); Gordon 1931: 527; Shen 1940a: 215; Davie 1992b: tab. 1; Wong et al. 2021: 17, fig. 28a, pl. 5F. Arcania novemspinosa — Hill 1982: 201, pl. 4D. Taxonomy: Shen (1931a), Sakai (1937), Naruse (2014). Type: Lectotype ♂ in RMNH D 790; and paratypes in RMNH (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: Western Indian Ocean to Central Pacific: from South Africa, Seychelles, India, to Southeast and East Asia, to northern Australia, Loyalty Islands and Marquesas Islands (Galil 2001a). Habitat: Subtidal; substrates of sandy-mud, muddy-sand or soft mud; depths 22 to 210 m (Chen and Sun 2002). Ebalia hayamaensis Sakai, 1963 Ebalia hayamaensis — Thompson and Shin 1983: tab. 4. Taxonomy: Sakai (1963), Komatsu (2011). Type: Holotype ♀ in BLIH. Type locality: Hayama, Sagami Bay, Japan. Distribution: Endemic to Japan: Sagami Bay and Ogasawara Islands, Japan (Takeda 1977; Komatsu 2011). Habitat: Subtidal; of depths 65 to 124 m (Komatsu 2011). Remarks: Ebalia hayamaensis Sakai, 1963 is known for certain only from Japan thus far (Komatsu 2011). The single local record was based on benthic grab sampling within the Victoria Harbour. We retain this record with much hesitation; the specimen in question needs to be located and re-examined to confirm this record. Ebalia scabriuscula Ortmann, 1892 Ebalia scabriuscula — Morton and Morton 1983: 157. Taxonomy: Sakai (1937: only the female specimen, according to Chen and Sun 2002), Chen (1989), Chen and Sun (2002). Type: Holotype ♀ MZS 645 (Komai 1999). Type locality: Sagami Bay, Japan. Distribution: South China Sea and East Asia: the Philippines, South and East China Seas, and Japan (Chen and Sun 2002). Habitat: Subtidal; substrates sandy, muddy, shelly; depths 50 to 202 m (Chen and Sun 2002; see Remarks page 39 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan below). Remarks: Many Ebalia species, including the present, inhabit deeper waters (holotype of E. scabriuscula Ortmann, 1892 from depths of 100 fathoms). The only local record of this species was recorded from intertidal habitats, according to Morton and Morton (1983), “… found in more silty sand beaches…” (p. 157). This record is probably incorrect and belongs to another species, but until the specimens can be re-examined, we retain it with doubt. Hiplyra platycheir (De Haan, 1841) # Philyra platycheira — Stimpson 1858c: 160; Stimpson 1907: 154; Gee 1926: 161 [list]; Shen 1940a: 216. Philyra platychira — Hill 1982: 199, pl. 3B; Davie 1992b: tab. 1; Blackmore and Rainbow 2000: app. 1. Hiplyra platycheir — Wong et al. 2021: 18, fig. 28b, c, pl. 6A. Taxonomy: Sakai (1937: as Philyra), Chen and Sun (2002: as Philyra), Galil (2009). Type: Lectotype ♂ RMNH D 793, and paralectotypes in RMNH (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: East Asia: South and East China Seas, Taiwan and Japan (Galil 2009). Habitat: Subtidal; substrate sandy; depths 17 to 150 m (Chen and Sun 2002). Ixa cylindrus (Fabricius, 1777) Ixa cylindrus — Shen 1940a: 216; Hill 1982: 201, pl. 4A. Taxonomy: Holthuis and Gottlieb (1956), Chen and Sun (2002), Takeda and Nagai (2009). Type: Syntypes in NHMD (Zimsen 1964). Type locality: Tharangambadi (= Tranquebar), India. Distribution: Western Indian Ocean to West Pacific: from East Africa, India, Singapore, Thailand, South China Sea, Taiwan, and Australia (Chen and Sun 2002; Davie 2002). Habitat: Subtidal; substrates of fine sand, muddy sand or soft mud; depths 19 to 33 m (Chen and Sun 2002). Ixa edwardsii Lucas, 1858 # Ixa cf. inermis — Hill 1982: 201, pl. 4B [not I. inermis Leach, 1817]. Ixa edwardsii — Wong et al. 2021: 19, fig. 29, pl. 6B. Taxonomy: Holthuis and Gottlieb (1956), Chen (1989), CGS Tan (1996), Chen and Sun (2002), Takeda and Nagai (2009). Type: Status unknown. Type locality: Unknown. Distribution: Western Indian Ocean to South China Sea: from East Africa, Red Sea, Persian/Arabian Gulf, India, to South China, Taiwan, Philippines and Indonesia (Holthuis and Gottlieb 1956; Chen and Sun 2002). Habitat: Subtidal; substrates of muddy-sand, sandy-mud or soft mud; depths 16 to 76 m (Chen and Sun 2002). Ixoides cornuta MacGilchrist, 1905 Ixoides coruntus (?) — Gordon 1931: 527, 530, text-fig. 7. Ixoides cornutus — Shen 1940a: 215. Taxonomy: Serène and Lohavanijaya (1973), Chen (1989), Galil (2001a: as Arcania), Chen and Sun (2002). Type: Syntypes probably in NHM (Davie 2002). Type locality: Persian Gulf. Distribution: Western Indian Ocean to Central Pacific: from Madagascar, East Africa, Persian/Arabian Gulf, to Southeast and East Asia, to New Caledonia, Vanuatu and Fiji (Galil 2001a). Habitat: Subtidal; substrates sandy-muddy; depths 28 to 204 m (Chen and Sun 2002). Remarks: The genus Ixiodes was synonymized under Arcania by Galil (2001) but Ng et al. (2017) argued that it should be recognized as a separate genus as it had several diagnostic morphological features. An ongoing phylogenetic study of these genera by Y.-J. Shih and B. Galil supports this. Lyphira ngankee Rahayu & Ng, 2024 # ? Philyra tuberculosa — Shen 1931b: 185, text-figs. 1–4 [not P. tuberculosa Stimpson, 1858 = P. carinata Bell, 1855]. ? Philyra globosa — Shen 1940a: 216 [not C. globosus Fabricius, 1793 = C. globus Fabricius, 1775]. Philyra cf. laminata — Hill 1982: 199, pl. 3C [not P. laminata Doflein, 1904]. Philyra heterograna — Blackmore and Rainbow 2000: app. 1. Lyphira heterograna — Wong et al. 2021: 20, fig. 30a–c, pl. 6C. Lyphira ngankee Rahayu & Ng, 2024: 376, figs. 1E–J, 5A, B, 6D, 8A, 9D, 10E, 11E, 13F, G, 14D, 15F, 16E, 18A–E, 20A–D, 21D. Taxonomy: Rahayu and Ng (2024). Type: Holotype ♂ ZRC 1999.0456, and paratypes in ZRC. Type locality: Nanao Island, Guangdong, China. Distribution: South China: so far only from Guangdong and Hong Kong. Habitat: Subtidal; substrate of soft mud or broken shells. Remarks: There are two primary issues associated with the taxonomy of this species. The first pertains to page 40 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan the identities of specimens initially referred to “Philyra tuberculosa” by Shen in 1931(b) from Hong Kong; and the second is with identities of L. heterograna sensu stricto. Shen (1931b) illustrated material he identified as “P. tuberculosa” from Hong Kong in considerable detail. He added more figures of the species (Shen 1932) when he compared it with his new species, Philyra peitaihoensis Shen, 1932, from northern China. Shen (1940a) subsequently decided that his Hong Kong material should be identified as “Philyra globosa” instead. Galil (2009) synonymized P. peitaihoensis as well as Phylira acutidens Chen, 1987, under Lyphira heterograna (Ortmann, 1892), figuring the overall male habitus and a G1 from southern Malaysia (not Indonesia as was stated). Previous records from China were referred to L. heterograna by Galil (2009). Philyra tuberculosa s. str. is now regarded as a junior synonym of P. carinata (Bell, 1855) (Galil 2009; see below). The name “Philyra globosa” presented in Shen (1940a) (citing Fabricius), can be traced back to the names Cancer globus Fabricius, 1775 and C. globosus Fabricius, 1793, but both represent the same species (Galil 2009). As shown by Galil (2009), the types of Cancer globus Fabricius, 1775 consist of two specimens, among which one smaller female was designated as the lectotype of that species by Holthuis (1962). The other male specimen, what H. Milne Edwards (1837) called “Philyra globulosa”, was shown to be P. heterograna (s. lato) instead. Rahayu and Ng (2024) recently revised the taxonomy of Lyphyira heterograna, examined and figured the types of L. heterograna (type locality: Japan), as well as of specimens from Japan, Korea, East and Southeast Asia, including Indonesian Papua, and showed that it is a complex of six species. Philyra peitaihoensis was confirmed to be synonymous with L. heterograna s. str., while P. acutidens was regarded as a valid Lyphira species and not just representing juveniles as had been supposed by Galil (2009). The specimens Galil (2009) figured as “P. heterograna” from Malaysia were shown to belong to new species. Rahayu and Ng (2024) referred material from Hong Kong and southern China, including the records of “P. tuberculosa” by Shen (1931b 1932) and “L. heterograna” by Wong et al. (2021), to a new species. This species, now L. ngankee, differs markedly from L. heterograna s. str. in possessing a distinct endostomial spine (see Shen 1931b: fig. 1; 1932: fig. 11c) (versus absent) and the exopod of the third maxilliped is proportionately much wider (see Shen 1931b: fig. 2; 1932: fig. 11f), characters already observed by Shen (1931b 1932) when he compared his material of P. peitaihoensis (= L. heterograna s. str.) with “P. tuberculosa” (= L. ngankee) from Hong Kong. There are also differences in the G1 as well as proportions of the chelae and ambulatory meri. Myra celeris Galil, 2001 # Myra fugax — Stimpson 1907: 152; Gee 1926: 161 [list]; Shen 1931a: 108, pl. 10(2); Gordon 1931: 527; Shen 1940a: 217; Hill 1982: 196, pl. 1A; Morton and Morton 1983: 187, fig. 10.4(3); Blackmore and Rainbow 2000: app. 1 [not Leucosia fugax Fabricius, 1798]. Myra sp. — Hill 1982: 198, pl. 1B. Myra celeris — Wong et al. 2021: 20, fig. 30d–f, pl. 6D. Taxonomy: Galil (2001b), Chen and Sun (2002: as M. fugax). Type: Holotype ♂ RMNH D 43202, and paratypes in RMNH, UMZC and NHM. Type locality: Japan, locality unspecified. Distribution: West Pacific: from South and East China, Taiwan, Japan, to Indonesia, Australia, and New Caledonia (Galil 2001b). Habitat: Subtidal; depths 4 to 52 m (Galil 2001b). Remarks: Myra celeris differs from M. fugax (Fabricius, 1798) sensu stricto by comparatively shorter fingers in relative to palm, and all past records of “M. fugax” from East Asia represent M. celeris instead (Galil 2001b). Myra hainanica Chen & Türkay, 2001 Myra affinis — Hill 1982: 198, pl. 2A [not M. affinis Bell, 1855]. Taxonomy: Chen and Türkay (2001), Chen and Sun (2002). Type locality: Off west coast of Hainan, China. Type: Holotype ♂ IOCAS K218B-40 and paratypes in IOASQ (now IOCAS) and SMF. Distribution: South China (Hainan and Hong Kong). Habitat: Subtidal; substrates of sandy mud or soft mud; depths 18 to 30 m (Chen and Sun 2002). Nursia plicata (Herbst, 1803) # Nursia plicata — Stimpson 1858c: 161; Stimpson 1907: 160; Alcock 1896: 180; Wong et al. 2021: 21, fig. 31a, b, pl. 6E. Nursia hardwickii — Gee 1926: 161 [list] [not N. hardwichii Leach, 1817 = Cancer lar Fabricius, 1793]. Nursia lar — Shen 1940a: 217 [no new specimens] [not C. lar Fabricius, 1793]. Taxonomy: Alcock (1896), Sakai (1965 1976), Campbell and Stephenson (1970), Serène and Soh (1976), Chen and Sun (2002: as N. lar), Naruse and Ng (2006). Type: Holotype ♂ ZMB HERBST 2198 (K Sakai page 41 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan sandy; depths 10 to 100 m (Dai and Yang 1991). Family Inachidae Macleay, 1838 Subfamily Inachinae Macleay, 1838 Achaeus japonicus (De Haan, 1839) Achæus japonicus — Stimpson 1857: 25; Stimpson 1907: 18. Achaeus japonicus — Gee 1926: 166 [list]; Shen 1940a: 218. Taxonomy: Sakai (1938). Type: Syntype RMNH D 42106 (only mouthparts; see Yamaguchi and Baba 1993, Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: East Asia: China (Guangdong to East China Sea) and Japan (Dai and Yang 1991). Habitat: Subtidal; substrates usually weedy and rocky, sometimes sandy or muddy; depths 20 to 100 m (Dai and Yang 1991). Achaeus lacertosus Stimpson, 1857 # Achaeus lacertosus — Wong et al. 2021: 26, fig. 42, pl. 8E. Taxonomy: Griffin and Yaldwyn (1965), Griffin (1970b), Sakai (1976), Dai and Chen (1992). Type: Presumably lost. Type locality: Port Jackson, New South Wales, Australia. Distribution: Western Indian Ocean to West Pacific: from South Africa, Persian/Arabian Gulf, India, to South China, Japan, towards Java Sea, Aru and Kai Islands to Australia (Griffin and Tranter 1986). Habitat: Subtidal; muddy to sandy substrates with shells or coral rubble (Davie 2002). Achaeus tuberculatus Miers, 1879 # Achaeus tuberculatus — Dudgeon 1982: tab. 1; Wong et al. 2021: 27, fig. 42a–d, pl. 8F. Taxonomy: Shen (1937a), Sakai (1938). Type: Syntypes probably in NHM. Type locality: 33°10'N, 129°12'E (near Nagasaki, Japan). Distribution: East Asia: coasts of China north to Bohai Seas, and Japan (Shen 1937a; Takeda and Miyake 1969b; Griffin and Tranter 1986). Habitat: Subtidal; substrates muddy, muddy-sandy or shelly; depths 30 to 200 m (Dai and Yang 1991). Camposcia retusa (Latreille, 1829) # Camposcia rectusa (sic) — Shen 1940a: 218. Camposcia retusa — Dudgeon 1982: tab. 1; Wong et al. 2021: 27, fig. 42e, f, pl. 9A. Taxonomy: Sakai (1938), Takeda (1973). Type: Probably in MNHN if still extant (Davie 2002). Type locality: Unknown. Distribution: Western Indian Ocean to West Pacific: from South Africa, Red Sea, to South China Sea, Taiwan and Japan, Palau to Australia (Griffin and Tranter 1986; Ng et al. 2001; Yang et al. 2008). Habitat: Subtidal, occasionally intertidal; substrate rocky or weedy; depths 1 to 30 m (Dai and Yang 1991). Family Majidae Samouelle, 1819 Subfamily Majinae Samouelle, 1819 Leptomithrax eldredgei Richer de Forges & Ng, 2015 Leptomithrax eldredgei Richer de Forges & Ng, 2015: 124, figs. 1, 2 [“off Hong Kong”]. Taxonomy: Richer de Forges and Ng (2015). Type: Holotype ♂ ZRC 1970.2.17.1, and paratypes in ZRC. Type locality: South China Sea near Hong Kong. Distribution: South China: so far known only from type locality. Habitat: Subtidal; on continental shelf. Micippa philyra (Herbst, 1803) # Micippa philyra — Gordon 1931: 529; Shen 1940a: 219; Horikoshi and Takeda 1982: tab. 1; Morton and Morton 1983: 47. Taxonomy: Sakai (1938), Buitendijk (1939). Type: Holotype ♀ ZMB HERBST 0077 (K Sakai 1999). Type locality: East Indies. Distribution: Western Indian Ocean to West Pacific: from Mauritius, Maldives, to Singapore, South China, Taiwan, Japan, to Indonesia and Australia (Buitendijk 1939). Habitat: Subtidal to intertidal; rocky lower intertidal zone (Yeh et al. 2006); at coral reef and rocky reef, associated with coral sponges, rubble and algae; depths up to 90 m (Davie 2002). Micippa platipes Rüppell, 1830 Micippa platipes — Dudgeon 1982: tab. 1. Taxonomy: Sakai (1938), Buitendijk (1939). Type: Status unknown (Davie 2002). Type locality: Red Sea. page 48 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Distribution: Western Indian Ocean to West Pacific: from South Africa, Red Sea, India, Southeast and East Asia, to Indonesia, Coral Sea and New Caledonia (Buitendijk 1939; Davie 2002; Yang et al. 2008). Habitat: Subtidal to intertidal; on and under encrusted rocks and dead coral in coral reefs and reef flats (Davie 2002); or beaches near the shoreline to depths of 20 m (Dai and Yang 1991). Micippa thalia (Herbst, 1803) # Micippa thalia — Alcock 1895: 251; Dudgeon 1982: tab. 1; Thompson and Horikoshi 1982: tab. 1; Davie 1992b: tab. 1. Taxonomy: Sakai (1938), Barnard (1950). Type locality: East Indies. Type: Holotype ♀ ZMB HERBST 0075 (K Sakai 1999). Distribution: Western Indian Ocean to West Pacific: from South Africa, Red Sea, India, to South China, Japan, towards Australia and New Caledonia (Dai and Yang 1991; Davie 2002). Habitat: Subtidal; at coral reef and rock reef, associated with broken shell, coral and sponge; or on muddy, muddy-sandy or shelly bottoms; depths 20 to 100 m (Dai and Yang 1991; Davie 2002). Ovimaja compressipes (Miers, 1879) Leptomithrax compressipes — Shen 1940a: 219. Taxonomy: Dai (1981: as Maja brevispinosis), Griffin and Tranter (1986), Ng and Richer de Forges (2015). Type: Holotype ♀ NHM 1860.15 (Griffin and Tranter 1986; Ng and Richer de Forges 2015). Type locality: Guangzhou (as Canton), China. Distribution: South China Sea and East Asia: from Indonesia, Thailand, the Philippines, to South China, Taiwan and Japan (Ng and Richer de Forges 2015). Habitat: Subtidal; substrates sandy-muddy; depths less than 50 m (S-H Wu 1999). Prismatopus aculeatus (H. Milne Edwards, 1834) Chlorinoides aculeatus — Blackmore and Rainbow 2000: app. 1. Taxonomy: Sakai (1938: as Acanthophrys), Griffin and Tranter (1986: as Thacanophrys); Ng et al. (2001). Type: Status unknown, probably in MNHN (Davie 2002). Type locality: “mers d’Asie”: seas of Asia. Distribution: Eastern Indian Ocean to West Pacific: from Sri Lanka, Mergui Archipelago, to Singapore, Thailand, South China and Taiwan, and Torres Strait and Australia (Sakai 1976; Yang et al. 2008). Habitat: Subtidal; substrates rocky and weedy (Dai and Yang 1991); body often decorated with corals and sponges (Davie 2002). Schizophrys aspera (H. Milne Edwards, 1834) # Schizophrys aspera — Shen 1940a: 219; Horikoshi and Takeda 1982: tab. 1; Dudgeon 1982: tab. 1; Morton and Morton 1983: 274. Taxonomy: Griffin and Tranter (1986), BY Lee et al. (2018). Type: Lectotype ♀ MNHN-B463 (designated by BY Lee et al. 2018). Type locality: Unknown. Distribution: Western Indian Ocean to West Pacific: from South Africa, Red Sea, to South China, Taiwan Japan, towards Fiji and Australia (Griffin and Tranter 1986; Ng et al. 2001; Yang et al. 2008); probably introduced to New Zealand (Ahyong and Wilkens 2011) and Hawaii (Carlton and Eldredge 2009; Castro 2011). Habitat: Subtidal to subtidal; rocky bottoms near to the shore, and sandy bottoms of coral reefs (Dai and Yang 1991). Remarks: B.Y. Lee et al. (2018) discussed the many taxonomic and nomenclatural problems with this species. Superfamily Orithyioidea Dana, 1852 Family Orithyiidae Dana, 1852 Orithyia sinica (Linnaeus, 1771) # Orithyia mammillaris — Shen 1931a: 106, pl. 9(1–3); Shen 1940a: 214. Orithyia sinica — Wong et al. 2021: 28, fig. 43, pl. 9B. Taxonomy: Chen (1993), Chen and Sun (2002), Ng et al. (2008). Type: Status unknown (Holm 1957; Wallin 2001). Type locality: China, locality unspecified. Distribution: East Asia: along coasts of China to Korea. Habitat: Subtidal; shallow seas of muddy-sandy substrates (Chen and Sun 2002). page 49 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Superfamily Palicoidea Bouvier, 1898 Family Palicidae Bouvier, 1898 Parapalicus trituberculatus (Chen, 1981) Palicus bidentatus Sakai, 1983a: 630, fig. 3g–j [southeast of Hong Kong Island, 21°42'N, 114°50'E]. Parapalicus trituberculatus — Castro 2000: 517, figs. 26c, 28, 58 [southeast of Hong Kong: 21°42'N, 114°50'E]. Taxonomy: Chen (1981: as Palicus), Sakai (1983a: as Palicus bidentatus), Castro (2000). Type: Holotype ♂ S209B-14, and paratypes in IOASQ (now IOCAS). Type locality: South China Sea southeast of Hong Kong (21°N, 114°30'E). Distribution: West Pacific: from Aru Sea, Indonesia, the Philippines, Gulf of Tonkin to northern South China Sea (Castro 2000). Habitat: Subtidal; on continental shelf from depths 69 to 122 m (Castro 2000). Superfamily Parthenopoidea Macleay, 1838 Family Parthenopidae Macleay, 1838 Subfamily Parthenopinae Macleay, 1838 Cryptopodia fornicata (Fabricius, 1787) # Cryptopodia fornicata — Stimpson 1857: 220; Stimpson 1907: 31; Gee 1926: 166 [list]; Shen 1940a: 217 [no new specimens]; Chiong and Ng 1998: 161, figs. 2–7, 10B; Wong et al. 2021: 29, fig. 44, pl. 9C. Taxonomy: Chiong and Ng (1998). Type: Lectotype ♀ ZMUC Cru 397 (now NHMD) (designated by Chiong and Ng 1998). Type locality: Tharangambadi (as Tranquebar), India. Distribution: Central Indian Ocean to South China Sea and East Asia: Central Indian Ocean, Andaman Islands, Singapore, the Philippines, Borneo, Indonesia, South China and Japan (Chiong and Ng 1998). Habitat: Subtidal, substrates broken-shelly and muddy-sandy; depths 25 to 30 m (Dai and Yang 1991). Cryptopodia transitans (Ortmann, 1893) Cryptopodia angusta Rathbun, 1916: 559 [“near Hong Kong”: 21°42'N, 114°50'E]. ? Heterocrypta transitans — Thompson and Horikoshi 1982: tab. 1. Taxonomy: Sakai (1965 1976: as Heterocrypta), SH Tan and Ng (2003). Type: Syntypes in MZS 920 (Komai 1999: as Heterocrypta). Type locality: Sagami Bay, Japan. Distribution: South China Sea and East Asia: the Philippines, South China, Taiwan and Japan (Sakai 1976). Habitat: Subtidal: substrates sandy or brokenshelly; depths 30 to 150 m (Sakai 1976). Enoplolambrus echinatus (Herbst, 1790) # Lambrus tuberculosus Stimpson, 1857: 220; Stimpson 1907: 29; Shen 1940a: 217. Parthenope tuberculosa — Gee 1926: 166 [list; “near Hong Kong”]. Parthenope tuberculosis — SY Lee et al. 2000: tab. 3. Enoplolambrus echinatus — Wong et al. 2021: 30, fig. 45, pl. 9D. Taxonomy: Flipse (1930: as L. (Platylambrus) echinatus and L. (P.) echinatus var. granulosus), SH Tan et al. (1999: as Platylambrus). Type: Lectotype ♀ ZMB HERBST 2055 and paralectotype in ZMB (K Sakai 1999). Type locality: Tharangambadi (as Tranquebar), India. Distribution: Eastern Indian Ocean to South China Sea: from East India, Singapore, Thailand, Vietnam, to South and East China, Taiwan, and Indonesia (Flipse 1930; SH Tan et al. 1999). Habitat: Subtidal, substrates broken-shelly or muddy; depths about 30 m deep (Dai and Yang 1991). Enoplolambrus laciniatus (De Haan, 1839) # Lambrus laciniatus — Stimpson 1857: 220; Stimpson 1907: 29. Lambrus validus — Shen 1931a: 109, pl. 10(3); Shen 1940a: 217; Morton and Morton 1983: 187, fig. 10.4(1) [not Parthenope (Lambrus) validus De Haan, 1837]. Enoplolambrus laciniatus — Wong et al. 2021: 30, fig. 46, pl. 9E, F. Taxonomy: Sakai (1976: as Parthenope (Platylambrus) validus forma laciniatus), SH Lee et al. (2016). Type: Lectotype ♂ RMNH D 39124, and paralectotypes in RMHN and USNM (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: East Asia: South China, Korea, and Japan (Sakai 1976; SH Lee et al. 2016; Wong et al. 2021). Habitat: Subtidal; substrates sandy or muddy, of shallow estuarine seas (material from local trawl surveys). Remarks: Based on material from recent local trawl surveys, three species of Enoplolambrus are present in seas adjacent to Hong Kong, of which E. laciniatus is the most common one. Past local records page 50 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan of Lambrus (or Parthenope) validus probably include E. laciniatus. Precise illustrations of local material were provided by Wong et al. (2021). Enoplolambrus validus (De Haan, 1837) # ? Lambrus validus — Balss 1922a: 134; Gordon 1931: 529; Thompson and Horikoshi 1982: tab. 1; Thompson and Shin 1983: tab. 4. ? Parthenope validus — Gee 1926: 166 [list]; Hill et al. 1978: 92, 114; Davie 1992b: tab. 1; Morton and Ruxton 1992: 1 unnumb. fig. (50); Blackmore and Rainbow 2000: app. 1; Shin 2000: tab. 1. ? Parthenope valida — Ong Che and Morton 1991: tab. 1. Enoplombrus validus — Wong et al. 2021: 31, fig. 47, pl. 10A. Taxonomy: Sakai (1976: as Parthenope (Platylambrus) validus forma validus). Type: Lectotype ♀ and paralectotypes in RMNH D 701, and paralectotypes in RMNH (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: West Pacific: China, Taiwan, Korea and Japan (Shen 1932b; Kim 1973; Sakai 1976; Dai and Yang 1991; Wong et al. 2021), likely also the Philippines (Estampador 1937), Indonesia (Buitendijk 1939) and Queensland, Australia (Campbell and Stephenson 1970). Habitat: Subtidal; substrates of sand, mud or broken shells; depths 30 to 200 m (Sakai 1976). Remarks: See Remarks under E. laciniatus above. Parthenope longimanus (Linnaeus, 1758) Lambrus longimanus — Gordon 1931: 529; Shen 1940a: 217. Taxonomy: Flipse (1930: as Lambrus (Lambrus)), SH Tan et al. (1999). Type: Status unknown (see below). Type locality: “Oceano Asiatico”: Asian Seas. Distribution: Western Indian Ocean to West Pacific: from Mauritius, India, Southeast and East Asia reaching Japan, to Indonesia, New Guinea and Australia (Flipse 1930; Sakai 1976; Davie 2002). Habitat: Subtidal, substrates muddy-sandy; depths 60 to 70 m (Dai and Yang 1991). Remarks: Extant Linnaean material compiled by Holm (1957: p. 56) and Wallin (2001: p. 16) includes one male individual of Cancer longimanus Linnaeus, 1758. However, this specimen was indicated to be a donation from “Gustav IV Adolf”, among material examined by Linnaeus after 1758 (Holm 1957; Wallin 1992) and thus is not a type of the species. Rhinolambrus lippus (Lanchester, 1902) # (Figs. 13, 36D, E) Rhinolambrus lippus — present record. Taxonomy: Lanchester (1902: as Lambrus), Trivedi and SH Tan (2019). Type: Holotype ♂ UMZCI 10519 (Trivedi and SH Tan 2019). Type locality: Peninsular Malaysia. Distribution: Western Indian Ocean to South China Sea: Madagascar, Djibouti, India and Peninsular Malaysia (Lanchester 1902; Nobili 1906; Trivedi and SH Tan 2019), to South China. Habitat: Shallow subtidal: substrate of rock fragments; depths around 5 m (local material). Remarks: Morphology of the only locally collected male agrees well with original descriptions based on material from an unknown locality in the Peninsular Malaysia (Lanchester 1902), extending distribution of this interesting species from the Indian Ocean to and northern South China Sea (Hong Kong). This species is a new record for the fauna of Hong Kong. Superfamily Pilumnoidea Samouelle, 1819 Family Galenidae Alcock, 1898 Subfamily Dentoxanthinae Števčić, 2005 Parapanope orientalis Ng & Guinot, 2021 # Parapanopeus euagora — SY Lee and Leung 1999: tab. 2 [not P. euagora De Man, 1895]. Parapanope euagora — Wong et al. 2021: 32, fig. 51, pl. 10E [not P. euagora De Man, 1895]. Taxonomy: Shen (1937a: as P. euagora), Guinot (1985: as P. sp.), Ng and Guinot (2021). Type: Holotype ♂ ZRC 2020.0404, and paratypes in ZRC. Type locality: Shacheng, Fujian, China. Distribution: East Asia: South China to Zhejiang, Korea and Japan (Ng and Guinot 2021). Habitat: Subtidal; substrates sandy and shelly, or in shallow waters on coral reefs, to depths of 35 m (Sakai 1976; Ng and Guinot 2021). Remarks: Guinot (1985) had doubts about specimens she had from China and Japan, which were referred to P. euagora De Man, 1895. Ng and Guinot (2021) revised the genus with fresh material from East Asia and transferred relevant records, including those from Japan to a new species, P. orientalis. Local material recently reported as P. euagora by Wong et al. (2021), having anterolateral lobes broadly triangular rather than laciniated with margins sinuous, and gastric regions moderately raised, are here identified page 51 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan as P. orientalis. As for the subfamily, Ng and Mitra (2021) argued that Parapanopinae Števčić, 2005 and Dentoxanthinae Števčić, 2005 were subjective synonyms and selected the latter name to have priority. Subfamily Galeninae Alcock, 1898 Galene bispinosa (Herbst, 1783) # Galene bispinosa — Alcock 1898: 136; Gee 1926: 163 [list]; Gordon 1931: 527; Shen 1940a: 227; SY Lee and Leung 1999: tab. 2; Wong et al. 2021: 31, fig. 48, pl. 10B. Taxonomy: Guinot (1969b), Ng (1998). Type: Lectotype ♂ ZMB HERBST 0170, and paralectotypes in ZMB (designated by K Sakai 1999). Type locality: “East Indies”. Distribution: Eastern Indian Ocean to West Pacific: from East India, Mergui Archipelago, Singapore, South China, Taiwan, to northern Australia (Dai and Yang 1991; Davie 2002). Habitat: Subtidal; substrates sandy and muddy, shallow waters (Dai and Yang 1991). Subfamily Halimedinae Alcock, 1898 Halimede fragifer (De Haan, 1835) # Halimede fragifer — Stimpson 1858a: 33; Stimpson 1907: 50; Gee 1926: 163 [list; “near Hong Kong”]; Shen 1940a: 225 [no new specimens]; Galil 2000: 326, fig. 1A, B; Wong et al. 2021: 31, fig. 49, pl. 10C. Fig. 13. Rhinolambrus lippus (Lanchester, 1902) (male, 36.0 × 33.8 mm, Clear Water Bay): A, carapace and cheliped; B, carapace, lateral view; C, frontal region and maxilliped 3, ventral view; D, right chela, ventral view; E, pleon and thoracic sternites; F, left G1, ventral view; G, left G2, ventral view. page 52 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan ? Halimede thurstoni — Gee 1926: 163 [list]. ? Halimede tyche — Gordon 1931: 528; Shen 1940a: 225 [not Cancer tyche Herbst, 1801]. Taxonomy: Galil (2000). Type: Holotype ♂ RMNH D 42183 (Yamaguchi and Baba 1993; Fransen et al. 1997; Galil 2000). Type locality: Japan, locality unspecified. Distribution: West Pacific: South China, Taiwan, Japan, and New Caledonia. Habitat: Subtidal; substrates muddy or sandymuddy; depths 20 to 50 m (Dai and Yang 1991). Halimede ochtodes (Herbst, 1783) # Polycremnus verrucifer Stimpson, 1858a: 33; Stimpson 1907: 49, pl. 6(1); Gee 1926: 163 [list; “near Hong Kong”]; AC Evans 1967: 406. Halimede ochtodes — Gee 1926: 163 [list]; Gordon 1931: 528; Shen 1940a: 225; Davie 1992b: tab. 1; Galil 2000: 328, fig. 2A, B; Wong et al. 2021: 31, fig. 50, pl. 10D. Taxonomy: Galil (2000). Type: Holotype ♀ ZMB HERBST 0144 (K Sakai 1999). Type locality: “East Indies”. Distribution: Arabian Sea to West Pacific: from Pakistan, India, Southeast and East Asia to South China, to Indonesia and Australia (Galil 2000; Davie 2002). Habitat: Subtidal; substrates muddy or muddysandy; depths 20 to 50 m (Dai and Yang 1991). Family Pilumnidae Samouelle, 1819 Subfamily Eumedoninae Dana, 1852 Ceratocarcinus frontodentata (Shen, Dai & Chen, 1982) Harrovia frontodentata — Blackmore and Rainbow 2000: app. 1. Taxonomy: Shen et al. (1982: as Harrovia), Chia and Ng (1998). Type: Holotype ♂ and paratype ♂ in IOASQ (now IOCAS). Type locality: Haimen, Guangdong, China. Distribution: So far only from South China (Chia and Ng 1998). Habitat: Subtidal; possibly symbiotic with crinoids as in congeneric C. longimanus White, 1847 (Castro et al. 1995). Ceratocarcinus trilobatus (Sakai, 1938) Ceratocarcinus trilobatus — Morton 1988: 20, pl. 2. Taxonomy: Sakai (1938: as Harrovia), Chia and Ng (1998). Type: Status and whereabouts unknown. Type locality: Shimoda, Japan. Distribution: East Asia: South China and Japan. Habitat: Subtidal; symbiotic with comasterids (Sakai 1938), or from Dendronethya, a soft coral (Dai and Yang 1991). Harrovia albolineata Adams & White, 1849 # Ceratocarcinus albolineatus — Stimpson 1857: 221; Stimpson 1907: 33; Shen 1940a: 218 [no new specimens]. Harrovia albolineata — Gee 1926: 166 [list]; Wong et al. 2021: 32, fig. 52. pl. 10F. Taxonomy: Chia et al. (1993), Chia and Ng (1998). Type: Lectotype ♂ BMNH 43.6 (now NHM), and paralectotype ♀ in NHM (designated by Chia et al. 1993). Type locality: Philippine Islands. Distribution: West Pacific: Sahul Shelf and Sunda Shelf (Chia and Ng 1998). Habitat: Subtidal; associated with crinoids; depths 10 to 140 m (Chia and Ng 1998). Permanotus purpureus (Gordon, 1934) Harrovia bituberculata — Blackmore and Rainbow 2000: app. 1. Taxonomy: Gordon (1934: as Harrovia), Chia and Ng (1998). Type: Holotype ♂ IRSNB IG 9223 (Chia and Ng 1998). Type locality: now Irian Jaya, Indonesia (as Sorong, New Guinea). Distribution: West Pacific: from Australia, Papua New Guinea, Indonesia, East Malaysia, to South China, Taiwan and Japan (Chia and Ng 1998; Yang et al. 2008). Habitat: Subtidal, of shallow waters, associated with crinoids (Chia and Ng 1998). Subfamily Pilumninae Samouelle, 1819 Actumnus dorsipes (Stimpson, 1858) Pilumnus dorsipes Stimpson, 1858a: 37; Stimpson 1907: 70, pl. 9(3, 3a); Gee 1926: 163 [list]. Actumnus asper — Shen 1940a: 227 [no new specimens]. Taxonomy: Sakai (1939), Takeda and Miyake (1969c). Type: Probably lost. Type locality: Hong Kong. Distribution: West Pacific: from northwestern Australia, the Philippines, to South China and Japan page 53 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan (Garth and Kim 1983; Dai and Yang 1991; Hosie et al. 2015). Habitat: Subtidal; substrates shelly; depths 20 to 60 m (Sakai 1976); or between crevices of rocks or coral reefs, or among roots of algae or within sponges at depths of 10 to 15 m (Dai and Yang 1991). Remarks: Shen (1940a), following Balss (1933), recognized A. dorsipes (Stimpson, 1858) a junior synonym of A. asper (Rüppell, 1830). Sakai (1939) examined material from Japan (A. dorsipes) and Palau (A. asper) and reported both as distinct, with material reported in Shen’s (1940a) and Stimpson’s (1858a) reports probably conspecific. Actumnus elegans De Man, 1888 Actumnus elegans — Clark et al. 2008: 921. Taxonomy: Chopra and Das (1937), Takeda and Miyake (1969c), Sakai (1983b). Type: Syntype RMNH D 1356c, other syntypes possibly also in ZMA (Fransen et al. 1997; Davie 2002). Type locality: Lampi Island (as Sullivan Island), Mergui Archipelago, Myanmar. Distribution: Eastern Indian Ocean to South China Sea and East Asia: Western Australia, Mergui Archipelago, the Philippines, South China, and Japan (Takeda and Miyake 1969c; Garth and Kim 1983; Davie 2002; Clark et al. 2008). Habitat: Subtidal; associated with epifauna, among coral head and sponges, often in areas with faster flowing waters (Goh et al. 1990; Clark et al. 2008). Remarks: The record presented by Clark et al. (2008) was identified based on work of Takeda and Miyake (1969c). The understanding of this species in the latter paper was considered erroneous by Sakai (1983b: p. 11), given frontal and anterolateral margins shown (Takeda and Miyake 1969c: fig. 1) differ substantially from the syntype in the ZSI, illustrated by Chopra and Das (1937: text-fig. 12). Actumnus setifer (De Haan, 1835) # Actumnus setifer — Alcock 1898: 202; Gordon 1931: 527; Shen 1940a: 227; Horikoshi and Takeda 1982: tab. 1; Johnson et al. 2008: 262, figs. 1–3; Wong et al. 2021: 33, fig. 53, pl. 11A. Taxonomy: Sakai (1939 1965), Takeda and Miyake (1969c), Campbell and Stephenson (1970). Type: Lectotype ♂ and paralectotype in RMNH D 42108, 42109, paralectotypes also in RMNH and MNHN (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: Western Indian Ocean to Central Pacific: from South Africa, Mauritius, Red Sea, to Southeast and East Asia reaching Japan, towards New Caledonia, Australia, Vanuatu, Fiji, Samoa, and Tahiti (Johnson et al. 2008). Habitat: Subtidal; in crevices of rock or coral reefs, or depths from littoral to 200 m (Dai and Yang 1991; Johnson et al. 2008); also reported to be excavating burrows in living or dead coral clumps (Johnson et al. 2008). Actumnus squamosus (De Haan, 1835) # Actumnus squamosus — Wong et al. 2021: 35, fig. 54, pl. 11B. Taxonomy: Sakai (1939 1965), Takeda and Miyake (1969c), Campbell and Stephenson (1970). Type: Holotype RMNH D 44480: only maxilliped 3 extant (Yamaguchi and Baba 1993; Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: Eastern Indian Ocean to West Pacific: from Palk Strait (India), Sri Lanka, Singapore, Southeast and East Asia reaching Japan, and Queensland (Australia) (Takeda and Miyake 1969c; Campbell and Stephenson 1970). Habitat: Intertidal to subtidal; to depths of 35 m (Sakai 1976). Aniptumnus quadridentatus (De Man, 1895) # (Figs. 14, 36F) Aniptumnus quadridentatus — present record. Taxonomy: Takeda (2001: as Pilumnopeus riui), Ng (2002b), Ng and Clark (2008), Hari et al. (2022). Type: Lectotype ♂ and paralectotype ♀ in RMNH D 1483 (designated by Ng 2002b; see also Fransen et al. 1997). Type locality: Pontianak, western Borneo, Indonesia. Distribution: South China Sea: so far from Papua (Indonesia), Borneo, Singapore, and South China (Takeda 2001; Ng and Clark 2008; Widyastuti and Rahayu 2022). Habitat: Intertidal; among barnacle clumps in mangroves (Mendoza and Ng 2011); locally shallow subtidal, proximal to the shore. Remarks: This species is a new record for the fauna of Hong Kong. Bathypilumnus sinensis (Gordon, 1930) Pilumnus sinensis Gordon, 1930: 523; Gordon 1931: 527, text-figs. 14, 15, 16(a, b); Shen 1940a: 228. page 54 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Taxonomy: Gordon (1931: as Pilumnus), Balss (1933: as Pilumnus), Ng and Tan (1984). Type: Holotype ♀ and paratype ♂ in NHM. Type locality: Hong Kong. Distribution: Arabian Sea to South China Sea: from Lakshadweep Islands (= Laccadives), Gulf of Thailand, to Singapore and South China (Ng and Tan 1984). Habitat: Subtidal; substrates sandy, shelly and muddy; depths 12 to 70 m (Ng and Tan 1984). Benthopanope eucratoides (Stimpson, 1858) # Heteropanope eucratoides Stimpson, 1858a: 35; Stimpson 1907: 64, pl. 8(2, 2a); Shen 1940a: 227. Heteropahope eucratoides — Gee 1926: 163 [list]. ? Heteropanope eucratoides — Melville and Morton 1983: 18, 1 unnumb. fig. (29). Taxonomy: De Man (1888b: as Heteropanope), Balss (1938: as Pilumnopeus), Davie (1989). Type: Presumably lost. Type locality: Hong Kong. Distribution: Eastern Indian Ocean to South China Sea: Mergui Archipelago, Malaysia and Singapore, and South China (Balss 1938; Lim et al. 1986). Habitat: Intertidal and shallow subtidal (Dai and Yang 1991). Glabropilumnus laevimanus (Dana, 1852) Pilumnus laevimanus — Gordon 1931: 527; Shen 1940a: 228 [no new specimens]. Glabropilumnus laevimanus — Galil and Takeda 1988: 72, figs. 3C, 4. Taxonomy: Galil and Takeda (1988). Type: Presumably lost. Type locality: Balabac Strait (as Balabac Passage), north of Borneo. Distribution: Western Indian Ocean to West Pacific: from East Africa, Red Sea, to Southeast and East Asia reaching Taiwan; Indonesia, Western Australia and New Caledonia (Galil and Takeda 1988). Habitat: Intertidal to shallow subtidal; coral or rocky reefs (Galil and Takeda 1988). Fig. 14. Aniptumnus quadridentatus (De Man, 1895) (male, 16.1 × 10.6 mm, off Castle Peak): A, carapace; B, major chela; C, P5, right; D, pleon; E, right G1, mesial view; F, right G1, lateral view. Structures A, C and D partially denuded; dashed line showing damage of specimen. page 55 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Glabropilumnus seminudus (Miers, 1884) Pilumnus seminudus — Gordon 1931: 542, text-fig. 17a. Glabropilumnus seminudus — Shen 1940a: 228. Taxonomy: Galil and Takeda (1988). Type locality: Thursday Island, Queensland, Australia. Type: Holotype ♀ BM 1882:7 and paratypes in NHM (Galil and Takeda 1988). Distribution: West Pacific: from northern Australia, Indonesia, to South China (Galil and Takeda 1988); exotic species in Hawaii (Carlton and Eldredge 2009). Habitat: Probably intertidal and shallow subtidal; recovered from fouling on hull of a barge (Edmondson 1962). Heteropanope glabra Stimpson, 1858 # Heteropanope glabra Stimpson, 1858a: 35; Stimpson 1907: 63, pl. 8(1); Gee 1926: 163 [list]; Shen 1940a: 227; J Jiang and Zhou 1982: 681. Taxonomy: Serène (1973a), Davie (1989). Type: Probably lost. Type locality: Hong Kong. Distribution: Western Indian Ocean to West Pacific: from East Africa, the Mergui Archipelago, Singapore, South China, Taiwan, to northern Australia and New Caledonia (Davie 1989). Habitat: Intertidal; in muddy and mangrove environments, sometimes with substantial freshwater influences, locally common among oyster clumps or rock crevices. Heteropilumnus ciliatus (Stimpson, 1858) Heteropilumnus ciliatus — Horikoshi and Takeda 1982: tab. 1; Morton and Morton 1983: 273, fig. 7.3(10). Taxonomy: Shen (1936a: as H. cristadentatus), Sakai (1939). Type: Presumably lost. Type locality: Shimoda (as Simoda), Japan. Distribution: East Asia: coasts of China to Shandong, Korea and Japan. Habitat: Intertidal; under rock fragments, on muddy substrates (Dai and Yang 1991). Heteropilumnus cristatus (Rathbun, 1909) # Heteropilumnus cristata — Wong et al. 2021: 36, fig. 55, pl. 11C. Taxonomy: Rathbun (1910: as Litochira). Type: Holotype ♀ and paratypes probably in NHMD (original text). Type locality: Two miles south of Koh Tutu, Gulf of Thailand. Distribution: South China Sea: so far from Gulf of Thailand and South China (Hong Kong). Habitat: Local female specimen collected among abandoned fish net, trawled from sandy-muddy substrates of depths 20 to 40 m. Heteropilumnus holthuisi Ng & Tan, 1988 # Litocheira subintegra — Gordon 1931: 528, text-fig. 25b. Heteropilumnus subinteger — Balss 1933: 42, pl. 7(32, 33); Shen 1936a: 65; Shen 1940a: 228. Taxonomy: Balss (1933: as H. subinteger), Ng and Tan (1988). Type: Holotype ♂ and paratypes in NHM 1900.10.22.175, and paratypes in ZRC. Type locality: Singapore. Distribution: Eastern Indian Ocean to South China Sea: from Mergui Islands to Singapore or Malacca, to South China (Guangxi to Fujian) (Ng and Tan 1988; Yang et al. 2008). Habitat: Intertidal and subtidal; under rocks on muddy substrata (Ng and Tan 1988). Remarks: As demonstrated in Ng and Tan (1988), the name Carcinoplax subinteger, established in Lanchester (1900: p. 750) was intended to “replace” the name Carcinoplax integra Miers, 1884, and thus based on the same type specimen. As such the name C. subinteger is not available. Carcinoplax integra s. str., a species from western Indian Ocean, is very distinct from the present West Pacific form previously called C. subinteger but referred to a new name, H. holthuisi by Ng and Tan (1988). Mier’s (1884) C. integra is now in Pseudolitochira Ward, 1942 (see Ng 1987; Maenosono 2019; Ng et al. 2021). Latopilumnus conicus Ng & Clark, 2008 Latopilumnus conicus Ng & Clark, 2008: 893, figs. 6–12; Clark et al. 2008: 922. Taxonomy: Ng and Clark (2008). Type locality: Conic Island, Hong Kong. Type: Holotype ♂ NHM 2003.690; paratypes in NHM and ZRC. Distribution: South China: so far from Hong Kong only. Habitat: Subtidal; within submarine caves (for description and discussions on sites, see Morton and Bamber 2008). page 56 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Nanopilumnus rouxi (Balss, 1935) # Nanopilumnus rouxi — Clark et al. 2008: 922. Taxonomy: Serène and Umali (1972: as Medaeus), Takeda (1974), Tirmizi and Ghani (1986). Type locality: Pamban, Gulf of Mannar, India. Type: Holotype ♂ in Basel Museum (now NMBA), Switzerland. Distribution: Arabian Sea to South China Sea and East Asia: from Pakistan, India, to Singapore, South China and Japan. Habitat: Shallow subtidal. Pilumnopeus convexus (Maccagno, 1936) # Pilumnopeus convexus — Wong et al. 2020: 2, figs. 1–4. Taxonomy: Davie (1989), Ghory et al. (2013), Wong et al. (2020). Type: Lectotype ♀ in MCG (designated by Davie 1989). Type locality: Assab, Eritrea (as Aseb, Ethiopia), southern Red Sea. Distribution: Western Indian Ocean: Chagos Archipelago, southern Red Sea, Persian/Arabian Gulf, Pakistan and India (Ghani and Davie 2000; Ghory et al. 2013); exotic species in Hong Kong (South China). Habitat: Intertidal and shallow subtidal; often among oyster clumps or fouling. Pilumnopeus makianus (Rathbun, 1931) # Pilumnopeus makiana — J Jiang and Zhou 1982: 681; ZG Huang and Lin 1993: app. Taxonomy: Shen (1932b: as Heteropanope), Balss (1933), Sakai (1939), Takeda and Miyake (1969c). Type: Holotype ♂ USNM 57497. Type locality: Shyago, Formosa (likely now Shehou, New Taipei City, Taiwan). Distribution: East Asia: coasts of China (East China to Bohai Seas), Taiwan, Korea and Japan (Takeda and Miyake 1969c; Yang et al. 2008). Habitat: Intertidal; under rocks on muddy or weedy substrates (Dai and Yang 1991). Pilumnus cursor A. Milne-Edwards, 1873 ? Pilumnus cursoni — J Jiang and Zhou 1982: 681. Taxonomy: Ng (1988a). Type: Syntype ♀ in MNHN (Ng 1988a). Type locality: New Caledonia. Distribution: Indo-West Pacific: so far confirmed from Phuket, Thailand and New Caledonia only (see Remarks below). Habitat: Subtidal; in coral rock (Ng and Davie 2002). Remarks: Records elsewhere previously recognized as “P. cursor” comprises a species complex, and various of these in the region had been referred to other species (Ng 1988a). This local record is thus very likely “either incorrect or suspicious” (Ng and Davie 2002: p. 382) and needs to be verified. Until then, we retain this record for the time being. Pilumnus longicornis Hilgendorf, 1878 # ? Pilumnus andersoni — Shen 1940a: 228. ? Pilumnus cf. andersoni — Horikoshi and Takeda 1982: tab. 1. Pilumnus longicornis — Horikoshi and Takeda 1982: tab. 1; Morton and Morton 1983: 273. Taxonomy: Balss (1933), Sakai (1939), Takeda and Miyake (1968). Type: Holotype ♂ in ZMB (Balss 1933). Type locality: Inhambane, Mozambique. Distribution: Western Indian Ocean to Central Pacific: from South Africa, Madagascar, Persian Gulf, Arabian Sea and Bay of Bengal, to Southeast and East Asia reaching Japan, and Indonesia, Australia, New Zealand towards Hawaii (Takeda and Miyake 1968; Garth and Kim 1983; Yang et al. 2008). Habitat: Intertidal and subtidal; rocky shores down depths of 10 to 20 m, also on sandy substrates down to depths of 85 m (Sakai 1976). Remarks: Balss (1933) synonymized P. andersoni De Man, 1887 (type locality Mergui Archipelago) under P. longicornis Hilgendorf, 1878, arguing that the morphological variations between the two (whether frontal and supraorbital margin denticulate) were agedependent. Shen (1940a), however, cited Alcock (1898: p. 191) who interpreted the two as distinct, listing the Hong Kong material as P. andersoni. Following Balss (1933) for the moment, we list the Hong Kong records under P. longicornis. Pilumnus minutus De Haan, 1835 # ? Pilumnus cf. minutus — Horikoshi and Takeda 1982: tab. 1; Wong et al. 2021: 36, fig. 56, pl. 11D. Pilumnus minutus — Ong Che and Morton 1992: tab. 1. Taxonomy: Sakai (1939), Takeda and Miyake (1968). Type: Status unknown (Davie 2002; not reported by Yamaguchi and Baba 1993). Type locality: Japan, locality unspecified. page 57 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Taxonomy: Yang and Tang (2006: as P. hastatides), Nguyen and Ng (2021: as Xiphonectes). Type: Holotype ♂ ZRC 2003.0527 and paratype ♀ in ZRC. Type locality: Teluk Tarempa, Anambas, Indonesia. Distribution: West Pacific: South China Sea to Taiwan and Japan, Indonesia and Australia (Nguyen and Ng 2021). Habitat: Subtidal; substrates of fine sand, muddy sand, soft mud or of shell fragments; depths 7 to 100 m (Yang et al. 2012). Remarks: Three Eodemus species (previously as Xiphonectes) are common in local shallow-water habitats, and at least four forms are sampled from local trawl surveys: X. dayawanensis (now E. unidens, see below), X. aff. hastatoides (yet described), X. pseudohastatoides, and “X. species” (Wong et al. 2021). These species were previously confounded under local records of “Portunus hastatoides” (Wong et al. 2010b 2021). Inspecting images of one of the syntypes of Portunus hastatoides Fabricius, 1798, Wong et al. (2010b) noted local material using this name might not be conspecific with the “real” X. hastatoides, and the local forms having the median frontal lobes less protruding than lateral lobes, comprise X. subtilis and “X. aff. hastatoides” (Wong et al. 2021). Recently, this issue was partially resolved by Nguyen and Ng (2021), who concluded X. hastatoides sensu stricto distributing from central Indian Ocean to Southeast Asia. The local form as “X. species”, distribution range of which spanning along the West Pacific, was described as X. subtilis (Nguyen and Ng 2021; now Eodemus). Eodemus unidens (Laurie, 1906) # ? Portunus hastatoides — Thompson and Horikoshi 1982: tab. 1 [part?]. ? Portunus trilobatus — Davie 1992b: tab. 1 [not P. (X.) trilobatus Stephenson, 1972]. Portunus dayawanensis — Wong et al. 2010b: 674, figs. 1E–I, 2C, D. Xiphonectes dayawanensis — Wong et al. 2021: 41, fig. 73, pl. 14C. Xiphonectes unidens — Nguyen and Ng 2021: 397, figs. 5D–H, 6G, H, 7. Taxonomy: Chen (1986: as Portunus dayawanensis), Wong et al. (2010b: as P. dayawanensis), Yang et al. (2012: as P. dayawanensis), Nguyen and Ng (2021: as Xiphonectes). Type: Holotype ♂ NHM 1907.5.22.309 (Nguyen and Ng 2021). Type locality: Gulf of Manaar, Sri Lanka. Distribution: Eastern Indian Ocean to South China Sea and East Asia: Sri Lanka, Malaysia, Singapore, Gulf of Thailand, to South China (Nguyen and Ng 2021). Habitat: Subtidal; substrates sandy or gravelly, or of shell fragments, depths to around 20 m (local records from Tolo Channel, where the species is common). Remarks: Davie (1992b) was correct in pointing out many past local records of “P. hastatoides” represented, or included the present species, a form bearing three frontal lobes instead of four, then listed as P. trilobatus Stephenson, 1972(b) (type locality Sumatra, Indonesia). This form was later recorded as P. dayawanensis Chen, 1986 (Wong et al. 2010b 2021), a species described from Daya Bay, Guangdong. Revising the Xiphonectes hastatoides species complex, Nguyen and Ng (2021) recently demonstrated several Indo-Pacific forms sharing a tri-lobed front, including Neptunus (Hellenus) tweediei Shen, 1937, P. trilobatus and P. dayawanensis, being junior synonyms of N. (H.) hastatoides var. unidens Laurie, 1906 (now as Eodemus unidens), a decision we herein follow. Incultus tuberculosus (A. Milne-Edwards, 1861) Neptunus (Hellenus) tuberculatus — Gordon 1931: 527 [not Achelous tuberculatus Stimpson, 1860]. Neptunus (Hellenus) tuberculosus — Shen 1940a: 220. Taxonomy: Crosnier (1962: as Portunus), Stephenson and Rees (1967: as Portunus), Apel and Spiridonov (1998: as Portunus), Yang et al. (2012: as Portunus), Koch et al. (2022). Type: Holotype ♂ MNHN B.6073 (Apel and Spiridonov 1998). Type locality: Hawaii Islands (as Iles Sandwich), yet probably erroneous (Castro 2011). Distribution: Western Indian Ocean to Central Pacific: from Madagascar, East Africa, Gulf of Aden, Gulf of Oman, Phuket (Thailand), South China Sea, to Palau, Indonesia, Australia, towards Hawaii (Stephenson 1972a; Apel and Spiridonov 1998); record from Hawaii remains doubtful (Castro 2011). Habitat: Subtidal; substrates of soft mud, sandy mud, muddy sand or shell fragments; depths 21 to 580 m (Yang et al. 2012). Remarks: Gordon’s (1931) record of “N. (H.) tuberculatus” citing “A. M.-Edw.” (contrasting Acheloüs tuberculatus Stimpson, 1860, an East Pacific species; see Garth and Stephenson (1966)), is probably erroneous, and actually belongs to the present species. Another record of this species from “China Sea, vicinity Hong Kong” at 21°53'N, 115°51'E, collection of the Albatross Philippine Expedition (1907–1910), was reported by Stephenson and Rees (1967: p. 52). Monomia argentata (A. Milne-Edwards, 1861) # Neptunus (Amphitrite) argentatus — Gordon 1931: 527; Shen 1940a: 220. page 64 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Portunus argentatus — Stephenson and Rees 1967: 16, fig. 2. Monomia argentatus — Wong et al. 2021: 39, fig. 67, pl. 13C. Taxonomy: Sakai (1939: as Portunus (Amphitrite)), J-F Huang and Yu (1997: as Portunus), Yang et al. (2012: as Portunus), Koch et al. (2017). Type: Lectotype ♂ NHM 1947.21 (designated by Koch et al. 2017). Type locality: Mouth of Lundu River, Borneo. Distribution: South China Sea to East Asia: Indonesia, Borneo (Malaysia), the Philippines, Vietnam, South China, Taiwan and Japan (Koch et al. 2017). Habitat: Subtidal; substrates of sand, muddy sand or soft mud; depths 10 to 219 m (Yang et al. 2012). Monomia haanii (Stimpson, 1858) # Neptunus (Amphitrite) gladiator — Shen 1940a: 220. Portunus gladiator — Morton 1979b: 136, tab. 7.1; Morton and Morton 1983: fig. 7.4(7). Portunus haani — Blackmore and Rainbow 2000: app. 1; SY Lee et al. 2000: tab. 3. Monomia haanii — Wong et al. 2021: 39, fig. 68, pl. 13D. Taxonomy: Sakai (1976: as Portunus (M.) gladiator), J-F Huang and Yu (1997: as Portunus), Ng et al. (2008: as Portunus), Yang et al. (2012: as P. gladiator), Windsor et al. (2019). Type: Lectotype ♂ RMNH D 379, and paralectotypes in RMNH ad NHM (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: Eastern Indian Ocean and West Pacific: Western Australia, South China Sea, Taiwan and Japan (Windsor et al. 2019). Habitat: Subtidal; substrate muddy with coarse and fine sands or shell fragments; depths 12 to 103 m (Yang et al. 2012). Remarks: The confusing taxonomy of Amphitrite haani Stimpson, 1858(a) and Portunus gladiator Fabricius, 1798 have been addressed by numerous authors, and revised in detail by Windsor et al. (2019). Monomia rubromarginatus (Lanchester, 1900) Neptunus (Amphitrite) rubro-marginatus — Shen 1937b: 104, textfig. 3. Taxonomy: Shen (1937b: as Neptunus (Amphitrite)), Stephenson and Campbell (1959: as Portunus). Type: Holotype ♀ NHM 1900.10.22.156 (NHM Data Portal). Type locality: Singapore. Distribution: West Pacific: the Philippines, Malay Archipelago and Singapore, Australia and New Caledonia (Davie 2002; Richer de Forges and Ng 2006). Habitat: Subtidal; substrates sandy, to depths of 38 m (Davie 2002). Remarks: In a treatment of Singapore swimming crabs, Shen (1937b) listed “South China Sea; Hong Kong; Malay Archipelago” as distribution of N. (A.) rubromarginatus (p. 104), yet this record was not cited in his later checklist of Hong Kong brachyuran fauna in 1940(a). Subsequent authors (e.g., Stephenson and Campbell 1959; Stephenson 1967 1972a) had cited Shen’s (1937b) report, including Hong Kong, or northern portion of South China Sea, within the distribution range of this Australian species. Extensive studies of brachyuran fauna in this region for the decades, however, have revealed no records of this species (Dai and Yang 1991; Yang et al. 2012). It is here retained in the Hong Kong fauna with doubt. Portunus pelagicus (Linnaeus, 1758) # Lupa pelagica — Stimpson 1858a: 38; Stimpson 1907: 76. Neptunus pelagicus — Heller 1865: 27 [part: material from East and Southeast Asia only]; Alcock 1899: 34 [part: material from Japan and Hong Kong only]. Portunus pelagicus — Gee 1926: 161 [list]; Hill et al. 1975: 82; Morton 1979a: fig. 6.16a; Morton and Morton 1983: 94, fig. 7.4(10); Thompson and Shin 1983: tab. 4; RSS Wu and Shin 1998: 108; SY Lee and Leung 1999: 78; B Chan and Leung 2002: 24, fig. 2C; Leung et al. 2004: 95, 2 unnumb. figs.; Tsang et al. 2020: 70, 2 unnumb. figs.; Wong et al. 2021: 40, fig. 69, pl. 13E. Portunus (Portunus) pelagicus — Shen 1934: 37, text-fig. 1. Neptunus (Neptunus) pelagicus — Shen 1940a: 220. Taxonomy: Sakai (1939: as Neptunus (Neptunus)), Holthuis (2004), JCY Lai et al. (2010). Type: Neotype ♂ ZRC 2007.0235 (designated by JCY Lai et al. 2010). Type locality: Pulau Tekong, Singapore (neotype). Distribution: West Pacific: northern Australia, Indonesia, the Philippines, Vietnam, South China, Taiwan and Japan; hybrid populations in Bay of Bengal and Andaman Sea (with P. reticulatus (Herbst, 1784)), and northern Australia (with P. armatus (A. MilneEdwards, 1861) (JCY Lai et al. 2010). Habitat: Intertidal, beneath seagrass or rocks, on muddy substrate or in shallow pools; or subtidal, substrates sandy, muddy, or of soft mud in estuaries; depths 5 to 47 m (Yang et al. 2012). Portunus sanguinolentus (Herbst, 1783) # Lupa sanguinolenta — Stimpson 1858a: 38; Stimpson 1907: 76. Neptunus (Neptunus) sanguinolentus — Balss 1922b: 106; Gordon 1931: 527; Shen 1940a: 220. page 65 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Portunus sanguinolentus — Gee 1926: 161 [list]; Stephenson and Rees 1967: 45, fig. 12a, b; Morton 1979a: fig. 6.15; Morton 1979b: 136, tab. 7.1; Thompson and Horikoshi 1982: tab. 1; RSS Wu 1982: tab. 1; Morton and Morton 1983: 94, fig. 7.4(9); Thompson and Shin 1983: tab. 4; Morton 1988: 24, pl. 4; Davie 1992b: tab. 1; RSS Wu and Shin 1998: 108; SY Lee and Leung 1999: 78; Blackmore and Rainbow 2000: app. 1; B Chan and Leung 2002: 24, fig. 2B; Leung et al. 2004: 97, 1 unnumb. fig.; Lui et al. 2007: 637; Wong et al. 2021: 40, fig. 70, pl. 13f. Portunus (Portunus) sanguinolentus — Shen 1932a: 34, text-figs. 2, 3, pl. 7a. Taxonomy: Sakai (1939: as Neptunus (Neptunus)), Stephenson and Campbell (1959), Apel and Spiridonov (1998), Yang et al. (2012). Type: Holotype ♂ ZMB 2099 (Apel and Spiridonov 1998; K Sakai 1999). Type locality: Tharangambadi (= Tranquebar), India (Apel and Spiridonov 1998). Distribution: Western Indian Ocean to Central Pacific: from South Africa, Madagascar, Red Sea, Gulf of Oman, through Southeast and East Asia to Australia, New Zealand and French Polynesia (Apel and Spiridonov 1998; Yang et al. 2012). Habitat: Intertidal to subtidal; substrates sandy, muddy or of soft mud; depths to 80 m or deeper (Yang et al. 2012). Portunus trituberculatus (Miers, 1876) # Portunus (Portunus) trituberculatus — Shen 1932a: 32, text-fig. 1, pl. 6. Neptunus (Neptunus) trituberculatus — Shen 1940a: 220. Portunus trituberculatus — Hill et al. 1975: 82; Hill et al. 1978: 114; Morton 1979b: 136, tab. 7.1; Davie 1992b: tab. 1; Leung et al. 2004: 99, 1 unnumb. fig.; Wong et al. 2021: 40, fig. 71, pl. 14A. Portunus trituberculata — Morton and Morton 1983: 94, fig. 7.4(8). Taxonomy: Sakai (1939: as Neptunus (Neptunus)), Stephenson and Rees (1967), Yang et al. (2012). Type: Holotype probably in NHM; paratype RMNH D 44303 (Fransen et al. 1997). Type locality: “Coasts of China or Japan”. Distribution: Eastern Indian Ocean to South China Sea and East Asia: Phuket (Thailand), Peninsular Malaysia, coasts of China, Taiwan and Japan (Stephenson 1972a; Ng and Davie 2002; Yang et al. 2012). Habitat: Subtidal; substrates of muddy sand, shell fragments or soft mud; depths 8 to 100 m (Yang et al. 2012). Subfamily Thalamitinae Paulson, 1875 Charybdis acuta (A. Milne-Edwards, 1869) Charybdis (Goniosoma) acuta — Shen 1934: 39, text-figs. 2–5; Shen 1940a: 221. Taxonomy: Shen (1934), Leene (1938), Yang et al. (2012). Type: Syntype probably in MNHN. Type locality: Japan, locality unspecified. Distribution: East Asia: South China, Taiwan, Korea and Japan (Yang et al. 2012). Habitat: Subtidal; among seagrass; depths 10 to 20 m (Yang et al. 2012). Charybdis affinis Dana, 1852 # Charybdis (Goniosoma) barneyi Gordon, 1930: 522; Gordon 1931: 527, 536, text-fig. 13 (a, b, b’); Shen 1934: 42, text-figs. 6–8; Shen 1940a: 221. Charybdis (Charybdis) affinis — Leene 1938: 35, figs. 8, 9. ? Charybdis (Goniosoma) gordoni — Shen 1940a: 221. Charybdis affinis — Davie 1992b: tab. 1; SY Lee and Leung 1999: tab. 2; Wong et al. 2021: 44, fig. 78, pl. 15B, C. Taxonomy: Shen (1937b), Leene (1938), Wee and Ng (1995), Yang et al. (2012). Type: Probably lost. Type locality: Singapore. Distribution: Eastern Indian Ocean to South China Sea: Bay of Bengal, Mergui Archipelago, Malaysia and Singapore, Sumatra, to South China and Taiwan (Yang et al. 2012). Habitat: Intertidal and subtidal; substrates of sand or muddy sand, from shallow depths (Yang et al. 2012). Charybdis anisodon (De Haan, 1850) # Charybdis anisodon — Stimpson 1858a: 39; Stimpson 1907: 80, pl. 12(1); Gee 1926: 161 [list; “near Hong Kong”]; Davie 1992b: tab. 1; Blackmore and Rainbow 2000: app. 1; Wong et al. 2021: 45, fig. 79, pl. 25D. Charybdis (Goniosoma) anisodon — Gordon 1931: 527; Shen 1932a: 43, text-figs. 9, 10, pl. 9b; Shen 1940a: 221. Taxonomy: Leene (1938), Crosnier (1962), Holthuis (1993), Wee and Ng (1995), Yang et al. (2012). Type: Lectotype ♂ RMNH D 44189, and paralectotypes in RMNH (designated by Holthuis 1993; see also Fransen et al. 1997). Type locality: Moluccas, Indonesia. Distribution: Western Indian Ocean to West Pacific: from Madagascar, Red Sea, to South China Sea and Japan, and northern Australia and New Caledonia (Wee and Ng 1995). Habitat: Subtidal; substrates of sandy mud or rock reefs; depths 5 to 30 m (Yang et al. 2012). page 66 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Charybdis annulata (Fabricius, 1798) # (Figs. 15, 37D) Charybdis annulata — present record. Taxonomy: Leene (1938), Wee and Ng (1995), Apel and Spiridonov (1998), Yang et al. (2012). Type: Syntypes in NHMD (Zimsen 1964), and possibly at ZMB (Apel and Spiridonov 1998). Type locality: “Oceano Indico”: possibly Tharangambadi (= Tranquebar), India (Fransen et al. 1997). Distribution: Western Indian Ocean to West Pacific: from South Africa, Madagascar, Pakistan and India, Southeast and East Asia to Japan (Apel and Spiridonov 1998). Habitat: Intertidal and shallow subtidal; among boulders along rocky reefs. Remarks: This species is a new record for the fauna of Hong Kong. Charybdis bimaculata (Miers, 1886) # Charybdis (Gonioneptunus) bimaculata — Shen 1940a: 222. Charybdis bimaculata — Morton and Morton 1983: fig. 7.4(5); Davie 1992b: tab. 1; Blackmore and Rainbow 2000: app. 1; SY Lee et al. 2000: tab. 3; Wong et al. 2021: 45, fig. 80, pl. 15E. Taxonomy: Shen (1932b), Leene (1938), Stephenson et al. (1957), Yang et al. (2012). Type: Syntypes probably in NHM. Type locality: Kobé, Japan. Distribution: Western Indian Ocean to West Pacific: Maldives, Lakshadweep (= Laccadive Islands), Bay of Bengal, Thailand, East Asia to Japan, and eastern Australia (Stephenson 1972a; Yang et al. 2012). Habitat: Subtidal; in shallow waters among seagrass, or on substrates of mud, sand, or shell fragments; depths 20 to 430 m (Yang et al. 2012). Charybdis brevispinosa Leene, 1937 # Charybdis brevispinosa — Wong et al. 2021: 45, fig. 81, pl. 15F. Taxonomy: Leene (1937 1938), Wee and Ng (1995), Yang et al. (2012). Type: Holotype ♂ and paratypes in ZMA. Type locality: Java Sea, Indonesia (6°13'S, 107°57'E). Distribution: West Pacific: Java Sea, Indonesia, to Singapore and Peninsular Malaysia, and South China (Yang et al. 2012). Habitat: Subtidal; substrates muddy sandy; depths around 30 m (Yang et al. 2012). Charybdis feriata (Linnaeus, 1758) # Charybdis crucifera — Stimpson 1858a: 39; Stimpson 1907: 80. Goniosoma cruciferum — Miers 1886: 191. Charybdis cruciata — Gee 1926: 161 [list]; Thompson and Horikoshi 1982: tab. 1; RSS Wu 1982: tab. 1; Morton and Morton 1983: 94, fig. 7.4(6). Charybdis (Goniosoma) cruciata — Shen 1932a: 38, text-fig. 6, pl. 8; Shen 1940a: 221. Charybdis feriatus — Hill et al. 1975: 82; Morton 1976: 103; Hill et al. 1978: 114; Morton 1979b: tab. 7.1, figs. 7.1, 7.5a; Morton and Morton 1983: 143; Davie 1992b: tab. 1; RSS Wu and Shin 1998: 108; SY Lee and Leung 1999: tab. 2; Blackmore and Rainbow 2000: app. 1; B Chan and Leung 2002: 24, fig. 2D; Lui et al. 2007: tab. 1; Wong et al. 2021: 45, fig. 82, pl. 16A. Charybdis feriata — Morton 1988: 28, pl. 6. Charybdis (Charybdis) feriatus — Leung et al. 2004: 89, 1 unnumb. fig. Taxonomy: Leene (1938: as C. cruciata), Holthuis (1962), Wee and Ng (1995), Apel and Spiridonov (1998). Type locality: Ambon, Indonesia (lectotype designated by Holthuis 1962). Type: Status and whereabouts unknown: now considered lost (Apel and Spiridonov 1998). Distribution: Western Indian Ocean to West Pacific: from South Africa, Madagascar, Gulf of Aden, Persian/Arabian Gulf, Arabian Sea, Bay of Bengal, to Southeast and East Asia to Japan, and Australia (see Apel and Spiridonov (1998)); exotic species in the Mediterranean (Abelló and Hispano 2006; Galil 2011). Habitat: Subtidal; substrates of sand or sandy mud, or in shallow rocky or coral reefs; depths 10 to 40 m, also reaching down to 118 m (Yang et al. 2012); juveniles found symbiotic under bells of jellyfish (Morton 1988). Remarks: The name Cancer feriata Linnaeus, 1758 (a name established making reference to a Cancer Marinus Lævis in Rumphius 1705), was resolved by Holthuis (1962), who designated the specimen figured in the work of G.E. Rumphius (1705: pl. 6P) as the lectotype (see also Holthuis 1959). The type material from Ambon, Indonesia, however, is believed no longer extant (Apel and Spiridonov 1998; Davie 2002). The species is now recognized as the senior synonym of Cancer sexdentatus Herbst, 1783, Ca. crucifer Fabricius, 1792 and Ca. cruciata Herbst, 1794. Charybdis granulata (De Haan, 1833) # Charybdis granulata — Stimpson 1858a: 39; Stimpson 1907: 82; Wong et al. 2021: 46, fig. 83, pl. 16B. Charybdis (Charybdis) natator — Leung et al. 2004: 93, 1 unnumb. fig. [not Cancer natator Herbst, 1794]. Taxonomy: Sakai (1976), Crosnier (1984b), Wee and Ng (1995), Yang et al. (2012). page 67 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Type: Lectotype ♂ RMNH D 468 and paralectotypes in RMNH (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: Western Indian Ocean to West Pacific: East Africa, East India, Singapore, East Asia to Japan, to eastern Australia (Crosnier 1984b; Wee and Ng 1995; Ng and Davie 2002; Ahyong and Lee 2005). Habitat: Subtidal; substrates rocky or sandy; depths 15 to 35 m (Yang et al. 2012). Charybdis hellerii (A. Milne-Edwards, 1867) # Charybdis (Goniosoma) merguiensis — Alcock 1899: 55; Gordon 1931: 527; Shen 1940a: 221. Charybdis hellerii — Davie 1992b: tab. 1; Blackmore and Rainbow 2000: app. 1; Wong et al. 2021: 47, fig. 84, pl. 16C. Taxonomy: Leene (1938), Wee and Ng (1995), Apel and Spiridonov (1998), Yang et al. (2012). Type: Lectotype ♂ MNHN B.761 S (designated by Apel and Spiridonov 1998; see also Dessouassi et al. 2019) Type locality: New Caledonia. Distribution: Western Indian Ocean to West Pacific: from Madagascar, East Africa, Red Sea, Persian Gulf, Arabian Gulf, Bay of Bengal, Southeast and East Asia to Japan, and Australia; alien species in the Mediterranean (Galil 2011), New Zealand (Ahyong and Wilkens 2011), Northwest and Southwest Atlantic (Dineen et al. 2001; Ruiz et al. 2011; Tavares 2011), West Africa (Dessouassi et al. 2019) and likely Hawaii (Edmondson 1954; Castro 2011). Habitat: Intertidal to subtidal; under rocks, or shallow pits with seagrasses, or within coral reefs (Yang et al. 2012). Charybdis hongkongensis Shen, 1934 # Charybdis (Goniohellenus) hongkongensis Shen, 1934: 46, text-figs. 11, 12; Shen 1940a: 222. Charybdis hongkongensis — Wong et al. 2021: 48, fig. 85, pl. 16D. Taxonomy: Shen (1934), Leene (1938), Wee and Ng (1995). Type: Holotype ♂ ZMFMIB No. 12585: status unknown. Type locality: Hong Kong. Distribution: West Pacific: Banda Sea and Sulawesi, Indonesia, to Peninsular Malaysia, Thailand, South China and Taiwan (Yang et al. 2012). Habitat: Subtidal; substrates sandy or sandymuddy; depths 30 to 400 m (Yang et al. 2012). Charybdis japonica (A. Milne-Edwards, 1861) # Charybdis sexdentata — Stimpson 1858a: 39; Stimpson 1907: 81. Charybdis japonica — Gee 1926: 161 [list; “near Hong Kong”]; Hill et al. 1975: 48; Morton 1976: 103; Hill et al. 1978: 92; Melville and Morton 1983: 18, 1 unnumb. fig. (19); Morton and Morton 1983: fig. 7.4(2); Davie 1992b: tab. 1; ZG Huang and Lin 1993: app.; SY Lee and Leung 1999: tab. 2; Wong et al. 2021: 48, fig. 86, pl. 16E. Taxonomy: Shen (1932b), Leene (1938), Yang et al. (2012). Type: Lectotype ♂ RMNH D 461; paralectotypes in RMNH, NSMT, KPMNH, MNHN, NHMD, MNHN, NHM and USNM (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: Western Indian Ocean to East Asia: Red Sea, west of Peninsular Malaysia, coasts of China, Korea and Japan (Leene 1937; Stephenson 1972a b; Yang et al. 2012); exotic species in the Mediterranean (Galil 2011) and New Zealand (Ahyong and Wilkens 2011). Habitat: Intertidal to subtidal; substrates of seagrass, mud, sand or rocks, to depths of 50 m (Yang et al. 2012). Charybdis lucifera (Fabricius, 1798) # Charybdis lucifera — B Chan and Leung 2002: 24, fig. 2F; Lui et al. 2007: tab. 1; Wong et al. 2021: 49, fig. 87, pl. 16F. Charybdis (Charybdis) lucifera — Leung et al. 2004: 91, 1 unnumb. fig. Taxonomy: Leene (1938), Wee and Ng (1995), Apel and Spiridonov (1998). Type: Syntypes in ZMK and NHMD (Zimsen 1964). Type locality: “In oceano indico”; probably Tharangambadi (= Tranquebar), India (Apel and Spiridonov 1998). Distribution: Western Indian Ocean to West Pacific: from Red Sea, Pakistan, India, Sri Lanka, Southeast and East Asia to Japan, and Australia (Apel and Spiridonov 1998). Habitat: Subtidal; substrates sandy-muddy or of rocky reefs; depths 5 to 100 m (J-F Huang and Yu 1997). Charybdis miles (De Haan, 1835) # Charybdis miles — Stimpson 1858a: 39; Stimpson 1907: 82; Gee 1926: 161 [list]; Stephenson and Rees 1968b: 92, figs. 1A, E, 2A, pl. 12A; Morton and Morton 1983: fig. 7.4(3); Wong et al. 2021: 50, fig. 88, pl. 17A. Charybdis (Goniosoma) miles — Gordon 1931: 527; Shen 1940a: 221. page 68 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Taxonomy: Leene (1938), Stephenson and Rees (1968b), Wee and Ng (1995), Apel and Spiridonov (1998), Yang et al. (2012). Type locality: Japan, locality unspecified. Type: Lectotype ♂ RMNH D 465, paralectotypes in RMNH, NHM and MNHN (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Distribution: Arabian Sea to West Pacific: from Gulf of Oman, India, Southeast and East Asia to Japan, and Australia (Yang et al. 2012). Habitat: Subtidal; substrates of sandy mud, muddy sand or soft mud; depths 10 to 160 m (Yang et al. 2012). Charybdis natator (Herbst, 1794) # Charybdis natator — Gee 1926: 161 [list; part?]; Blackmore and Rainbow 2000: app. 1; Lui et al. 2007: tab 1; Wong et al. 2021: 51, fig. 89, pl. 17B. Charybdis (Goniosoma) natator — Shen 1932a: 40, text-figs. 7, 8, pl. 9a; Shen 1940a: 221. Taxonomy: Sakai (1976), Wee and Ng (1995), Apel and Spirodonov (1998), Yang et al. (2012). Type locality: “Das Veterland ist Ostindien”, could be east coast of India or the Indo-Malayan Archipelago (Apel and Spiridonov 1998). Type: Status unknown (not found in ZMB by Apel and Spiridonov 1998 by K Sakai 1999). Distribution: Western Indian Ocean to West Pacific: from South Africa, Madagascar, Red Sea, Persian Gulf, Arabian Sea, Bay of Bengal, Southeast and East Asia to Japan, and Australia (Apel and Spiridonov 1998). Habitat: Subtidal; substrates of sand or sandy mud; depths 30 to 310 m (Yang et al. 2012). Remarks: Past local records of C. natator (since Gee 1926) might include C. granulata (De Haan, 1833). The latter species was long considered a junior synonym of C. natator until Sakai (1976) showed both being distinct (see also Wee and Ng 1995). Charybdis orientalis Dana, 1852 # (Figs. 16, 37E) Charybdis orientalis — present record. Taxonomy: Leene (1938), Stephenson et al. (1957), Wee and Ng (1995), Apel and Spiridonov (1998), Yang et al. (2012). Type: Syntype in MCZ (Apel and Spiridonov 1998). Type locality: Caldera Bay, Mindanao, the Philippines. Distribution: Western Indian Ocean to Central Pacific: from South Africa, Madagascar, Red Sea, Arabian Sea and Bay of Bengal, Southeast and East Asia to Japan, and Australia, probably also French Polynesia (Apel and Spiridonov 1998; Yang et al. 2012). Habitat: Subtidal; rocky substrates of sand, pebbles or with aquatic vegetation; depths 10 to 30 m (Yang et al. 2012). Remarks: This species is a new record for the fauna of Hong Kong. Charybdis truncata (Fabricius, 1798) # Charybdis truncata — Stimpson 1858a: 39; Stimpson 1907: 82; Shin and Thompson 1982: app.; Morton and Morton 1983: fig. 7.4(4); Davie 1992b: tab. 1; Mackie et al. 1993: app.; Blackmore and Rainbow 2000: app. 1.; SY Lee et al. 2000: tab. 3.; B Chan and Leung 2002: 24; Wong et al. 2021: 52, fig. 90, pl. 17C. Charybdis (Goniohellenus) ornata — Alcock 1899: 64. Charybdis (Goniohellenus) truncatus — Balss 1922b: 103; Gordon 1931: 527; Shen 1934: 49, text-figs. 13, 14. Charybdis (Goniohellenus) truncata — Shen 1940a: 222. Taxonomy: Shen (1934), Leene (1938), Crosnier (1962), Wee and Ng (1995), Yang et al. (2012). Type: Syntypes in NHMD (Zimsen 1964; see also Rathbun 1902). Type locality: “Habitat in Oceano Indico”, probably Tharangambadi (= Tranquebar), India (Fransen et al. 1997). Distribution: Western Indian Ocean to West Pacific: from Madagascar, Maldives, India, Sri Lanka, Myanmar, Southeast and East Asia, and Australia (Yang et al. 2012). Habitat: Subtidal; substrates of sandy-mud, soft mud, muddy sand or coarse sand; depths 7 to 107 m (Yang et al. 2012). Charybdis vadorum Alcock, 1899 # Charybdis (Goniohellenus) sinensis Gordon 1930: 522; Gordon 1931: 534, text-figs. 11, 12(c, d, d’); Shen 1934: 44, text-figs. 9, 10. Charybdis (Goniohellenus) vadorum — Shen 1940a: 222. Charybdis vadorum — Shin and Thompson 1982: app.; Thompson and Horikoshi 1982: tab. 1; RSS Wu 1982: tab. 1; Shin 1990: tab. 2; Ong Che and Morton 1991: tab. 1; Davie 1992b: tab. 1; RSS Wu and Shin 1998: 112; SY Lee and Leung 1999: tab. 2; Blackmore and Rainbow 2000: app. 1; SY Lee et al. 2000: tab. 3; Wong et al. 2021: 53, fig. 91, pl. 17D. Taxonomy: Chopra (1935), Leene (1938), Wee and Ng (1995), Apel and Spiridonov (1998), Yang et al. (2012). Type: Lectotype ♂ in ZSI, which G1 figured in text-figure 13 c, d in Chopra (1935) (designated by Apel and Spiridonov 1998). Type locality: Indo-Burmese coast of the Bay of Bengal. Distribution: Eastern Indian Ocean to South China page 69 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Sea: Bay of Bengal, Southeast and East Asia to South China (Apel and Spiridonov 1998). Habitat: Subtidal; substrates of sandy mud, soft mud, muddy sand or coarse sand; depths 10 to 90 m (Yang et al. 2012). Charybdis variegata (Fabricius, 1798) # Charybdis (Goniosoma) variegata — Alcock 1899: 60; Gordon 1931: 527; Shen 1940a: 222. Charybdis variegata — Balss 1922b: 104; Gee 1926: 161 [list]; Davie 1992b: tab. 1; SY Lee and Leung 1999: tab. 2; Blackmore and Rainbow 2000: app. 1; SY Lee et al. 2000: tab. 3; Wong et al. 2021: 94, fig. 92, pl. 17E. Taxonomy: Shen (1937b), Leene (1938), Wee and Ng (1995), Apel and Spiridonov (1998), Yang et al. (2012). Type: Lectotype ♂ and paralectotypes in NHMD (designated by Apel and Spiridonov 1998). Type locality: “In oceano indico”; probably Tharangambadi (= Tranquebar), India (Apel and Spiridonov 1998). Distribution: Western Indian Ocean to West Pacific: from South Africa, Madagascar, Red Sea, Pakistan, India, Southeast and East Asia to Japan, and northern Australia (Yang et al. 2012). Habitat: Subtidal; substrates muddy or sandy; depths 10 to 80 m (Yang et al. 2012). Goniosupradens acutifrons (De Man, 1879) # (Figs. 17, 37F) Charybdis cf. miles — Morton and Ruxton 1992: 50, 1 unnumb. fig.; Morton and Harper 1995: 46, pl. 24B; Morton and Ruxton 1997: 51, 1 unnumb. fig. [not Portunus (C.) miles De Haan, 1835]. Goniosupradens acutifrons — present record. Taxonomy: Leene (1936 1938), Crosnier and Thomassin (1974), Yang et al. (2012). Type: Holotype ♂ RMNH D 448 (Fransen et al. 1997). Type locality: Timor, Indonesia. Distribution: Western Indian Ocean to West Pacific: from Madagascar, Tanzania, Phuket (Thailand), to Southeast and East Asia to Japan, and Solomon Islands (Yang et al. 2012). Habitat: Subtidal; rocky or coral reefs; depths 5 to 30 m (Yang et al. 2012). Remarks: This species is a new record for the fauna of Hong Kong. Lissocarcinus arkati Kemp, 1923 Lissocarcinus arkati — Gordon 1931: 527, 533; Shen 1940a: 219. Taxonomy: Chopra (1935), Sakai (1939), Crosnier (1962), Yang et al. (2012). Type: Syntype ♀♀ ZSI C 693/1 (original text). Type locality: Sandheads off mouth of River Hooghly (as Hughli River), western Bengal, India. Distribution: Western Indian Ocean to West Pacific: from Madagascar, Bay of Bengal, to Southeast and East Asia to Japan, and Queensland (Australia). Habitat: Subtidal; substrates sandy-mud, soft mud or broken shells; depths 10 to 67 m (Yang et al. 2012). Lissocarcinus orbicularis Dana, 1852 Lissocarcinus orbicularis — Markham 1982: 357; Markham 1992: tab. 1 [list]; Li 2003: 153 [no new specimens]. Taxonomy: Sakai (1939), Crosnier (1962), Spiridonov (1999), Yang et al. (2012). Type: Presumably lost. Type locality: Ovalau, Fiji. Distribution: Western Indian Ocean to Central Pacific: South Africa, Madagascar, Red Sea, India and Sri Lanka, to Southeast and East Asia to Japan, and Australia, New Caledonia, Fiji, Hawaii and Tuamotu. Habitat: Subtidal; commensal with holothurian Holothuria (Halodeima) atra Jaeger, 1833, H. (Stauropora) pervicax Selenka, 1867 and Bohadschia argus Jaeger, 1833, in respiratory tree or cloaca of hosts, on various substrates (Spiridonov 1999). Thalamita chaptalii (Audouin, 1826) Thalamita chaptalii — Shen 1940a: 223. Taxonomy: Stephenson and Hudson (1957), Crosnier (1962), Wee and Ng (1995), Yang et al. (2012). Type: Status unknown. Type locality: Red Sea. Distribution: Western Indian Ocean to Central Pacific: from Madagascar, Red Sea, India, Sri Lanka, Southeast and East Asia to Taiwan, to Australia, New Caledonia, Solomon Islands and Tahiti (Stephenson 1972a; Yang et al. 2012). Habitat: Subtidal; among crevices of coral reefs or beneath rocks; depths 5 to 40 m (Yang et al. 2012). Thalamita imparimana Alcock, 1899 Thalamita imparimana — Stephenson and Rees 1967: 78, fig. 28, pl. 7A [China Sea, vicinity Hong Kong, 21°46'N, 114°47'E]. Taxonomy: Stephenson and Rees (1967), Serène and Soh (1976: as T. muusi), Yang et al. (2012). Type: Syntypes probably in ZSI. page 70 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Type locality: Ganjam Coast, India. Distribution: Eastern Indian Ocean to South China Sea: Northeastern India, Phuket (Thailand), the Philippines, South China. Habitat: Subtidal; on substrates sandy or beneath coral reefs; depths 53 to 69 m (Yang et al. 2012). Thalamita picta Stimpson, 1858 Thalamita picta — Thompson and Horikoshi 1982: tab. 1; Thompson et al. 1982: tab. 1; RSS Wu 1982: tab. 1; Morton and Morton 1983: 94, fig. 7.4(1); RSS Wu and Shin 1998: tab. 1. Taxonomy: Stephenson and Hudson (1957), Figs. 15–17. Charybdis annulata (Fabricius, 1798) (male, 80.3 × 54.1 mm, Shui Hau): A, carapace and cheliped; B, right chela. 16, Charybdis orientalis Dana, 1852 (male, 59.2 × 35.7 mm, Shelter Island): carapace and cheliped. 17, Goniosupradens acutifrons (De Man, 1879) (male, 81.8 × 52.2 mm, Bluff Island): carapace and cheliped. page 71 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Crosnier (1962), Garth (1965), Chen (1975), Wee and Ng (1995), Yang et al. (2012). Type: Presumably lost. Type locality: Oshima (as Ousima), Japan. Distribution: Western Indian Ocean to East Pacific: from East Africa, Red Sea, India, Southeast and East Asia to Japan, towards Australia, New Caledonia, Marshall Islands, Hawaii, Tuamotu, Samoa and Clipperton Island. Habitat: Intertidal and subtidal; in rocky or coral reefs (Yang et al. 2012). Remarks: This species is commonly reported associated with corals reefs (see cited works under Taxonomy above). Considering the habitat, some local records appear in need of verification: Thompson and Horikoshi (1982), Thompson et al. (1982), RSS Wu (1982) and RSS Wu and Shin (1998) all reporting material of this species sampled from Tolo Channel (trawled from gravelly benthic substrates). Thalamita sima H. Milne Edwards, 1834 # Thalamita sima — Stimpson 1858a: 39; Alcock 1899: 81; Stimpson 1907: 83, pl. 11(2); Gee 1926: 161 [list; “near Hong Kong”]; Gordon 1931: 527; Shen 1934: 54, text-figs. 17, 18; Shen 1940a: 223; Davie 1992b: tab. 1; Mackie et al. 1993: app.; Blackmore and Rainbow 2000: app. 1; Wong et al. 2021: 56, fig. 94, pl. 18A. Taxonomy: Stephenson and Hudson (1957), Wee and Ng (1995), Apel and Spiridonov (1998), Yang et al. (2012). Type: Probably in MNHN, status unknown (Apel and Spiridonov 1998). Type locality: Coromandel Coast, Southeast India. Distribution: Western Indian Ocean to Central Pacific: from East Africa, Red Sea, India, Sri Lanka, Southeast and East Asia to Japan, Australia, New Caledonia, New Zealand and Hawaii (Yang et al. 2012). Habitat: Intertidal and subtidal; on sandflats, substrates muddy, or muddy-sandy, to depths of 50 m (Yang et al. 2012). Thranita crenata (Rüppell, 1830) # Thalamita crenata — Stimpson 1907: 84, pl. 10(6, 6a); Gordon 1931: 527; Shen 1940a: 223; Davie 1992b: tab. 1; BKK Chan and Caley 2003: 86, 2 unnumb. figs.; Wong et al. 2021: 55, fig. 93, pl. 17F. Thranita crenata — Bravo et al. 2021: tab. 2. Taxonomy: Stephenson and Hudson (1957: as Thalamita), Crosnier (1962: as Thalamita), Wee and Ng (1995: as Thalamita), Yang et al. (2012: as Thalamita). Type locality: Southern Red Sea. Type: Lectotype ♂ SMF 5611, and paratypes in SMF and RMNH (designated by Apel and Spiridonov 1998). Distribution: Western Indian Ocean to Central Pacific: from South Africa, Madagascar, Red Sea, Gulf of Oman, Pakistan, India, Southeast and East Asia to Japan, to Australia, New Caledonia, Tonga, Tuamotu, and Hawaii. Habitat: Intertidal and subtidal; locally not uncommon beneath rocks on gravelly sandflats. Thranita danae (Stimpson, 1858) # Thalamita danae — Stimpson 1858a: 39; Stimpson 1907: 85, pl. 11(1, 1a); Gee 1926: 161 [list]; Shen 1934: 52, text-figs. 15, 16; Shen 1940a: 222; Morton and Harper 1995: 43, fig. 6(34); Wee and Ng 1995: 73, figs. 37A–C, 38A–C, 39A–C, 40A, B, 41A–D, 42A–I; Morton and Ruxton 1997: 1 unnumb. fig. (22); VCS Lai et al. 2006: 65, 3 unnumb. figs. ? Thalamita stimpsoni — Alcock 1899: 80. ? Scylla serrata — Morton 1979a: fig. 6.20g. Taxonomy: Stephenson and Hudson (1957: as Thalamita), Stephenson (1972b: as Thalamita), Wee and Ng (1995: as Thalamita), Yang et al. (2012: as Thalamita). Type: Neotype ♂ NHM 1986.848 (designated by Wee and Ng 1995). Type locality: Tolo Harbour, Hong Kong (neotype). Distribution: Western Indian Ocean to West Pacific: from East Africa, Red Sea, India, Southeastand East Asia to Japan, Australia, New Caledonia, Marshall Islands, Fiji and Samoa. Habitat: Intertidal and subtidal; among rocky reefs, or on substrates of sand or muddy sand. Remarks: While the species T. danae Stimpson, 1858 and T. stimpsoni A. Milne-Edwards, 1871 are merely distinguished by relative size of the fourth anterolateral tooth (Stephenson and Hudson 1957), the character is regarded as age-dependent, and the decision of synonymizing T. stimpsoni under T. danae as presented in Stephenson (1972b) and Wee and Ng (1995) is followed. Another closely similar species, T. foresti Crosnier, 1962, was reported by Stephenson and Rees (1967: p. 75) based on material from “Sabtan I., China Sea, vicinity Hong Kong” collected during the Albatross Philippine Expedition in 1908. The site was probably now Sabtang of the Batanes Islands, recorded to be around 20°46'N, 120°52'E (US Bureau of Fisheries 1910) within the Luzon Strait, therefore cannot be considered as fauna of Hong Kong. page 72 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Thranita pelsarti (Montgomery, 1931) # (Figs. 18, 37G) ? Thalamita danae — Morton and Harper 1995: 46, pl. 24C [not T. danae Stimpson, 1858]. Thalamita pelsarti — Tsang et al. 2020: 72, 2 unnumb. figs. Taxonomy: Wee and Ng (1995: as Thalamita), Crosnier (2002: as Thalamita), Yang et al. (2012: as Thalamita). Type: Syntypes in NHM 1931.7.24.41-42 (Crosnier 2002). Type locality: Pelsaert (= Pelsart) and Long Islands, Abrolhos Islands, Australia. Distribution: Eastern Indian Ocean to South China Sea and East Asia: verified from Abrolho Islands, Australia, Indonesia, Malaysia and Singapore, Vietnam, South China, Taiwan and Japan (Crosnier 2002; Yang et al. 2012). Habitat: Intertidal and subtidal; close to shores, on substrates muddy or rocky, also common among coral reefs (Yang et al. 2012). Remarks: This species had long been considered synonymous under Cancer prymna Herbst, 1803, until resurrected by Wee and Ng (1995) (see also Crosnier (2002)). Appearing rather common among coral communities in Hong Kong, very likely T. pelsarti has been confounded under past local records of T. prymna. Morphological distinctions between T. pelsarti and T. prymna s. str. have been elaborated in detail by Wee and Ng (1995) and recently by Y.-H. Huang and Shih (2021). The latter work showed, however, molecular evidences based on the mitochondrial COI marker showing substantial but intraspecific level of divergences differences, and both species as well as T. tenuipes (Borradaile, 1902) constitute a thus far unresolved species complex (Y-H Huang and Shih 2021). We herein list both taxa as distinct species. Fig. 18. Thranita pelsarti (Montgomery, 1931) (male, 54.8 × 33.6 mm, Sharp Island): A, carapace; B, basal antennal segment; C, left chela, external view; D, left chela, internal view; E, left chela, ventral view; F, left P5; G, left G1, mesial view; H, tip of left G1, mesial view. Structure A partially denuded. page 73 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Type locality: Djeddah, Red Sea. Distribution: Western Indian Ocean to Central Pacific: from South Africa, Madagascar, Red Sea, Bay of Bengal, South China Sea and South China to Indonesia and Australia, Palau, Tonga, Fiji, Samoa, Tahiti and Hawaii (Serène 1984). Habitat: Intertidal and subtidal; rocky or coral reefs (Dai and Yang 1991). Pilodius granulatus Stimpson, 1858 # Pilodius granulatus Stimpson, 1858a: 32; Stimpson 1907: 58, pl. 7(2); Gee 1926: 163 [list]; Morton 1976: 98; Morton 1979a: 113; Horikoshi and Takeda 1982: tab. 1; Clark and Galil 1993: 1133, figs. 5A–G, 33A, 41B. Taxonomy: Clark and Galil (1993). Type: Presumably lost. Type locality: Hong Kong. Distribution: West Pacific: northeastern Australia, Indonesia, Palau, Singapore, the Philippines, to South China (Clark and Galil 1993). Habitat: Subtidal; associated with coral reefs (Davie 2002). Pilodius miersi (Ward, 1936) # (Figs. 22, 38E) Pilodius miersi — present record. Taxonomy: Clark and Galil (1993), SH Lee and Ko (2011). Type: Holotype ♂ QM W745, and paratypes in NHM and AM (Davie 2002). Type locality: Lindeman Islands, Queensland, Australia. Distribution: West Pacific: northeastern Australia, Singapore, Vietnam, South China, Korea and Japan (Clark and Galil 1993; SH Lee and Ko 2011). Habitat: Subtidal; locally on coarse sandy substrates under boulders near coral communities. Remarks: This species is a new record for the fauna of Hong Kong. Fig. 21. Novactaea pulchella (A. Milne-Edwards, 1865) (male, 15.1 × 10.4 mm, Starfish Bay): A, carapace; B, maxilliped 3; C, right chela; D, right P5; E, thoracic sternites and pleon; F, left G1, mesial view; G, tip of left G1, mesial view. Structures A, C to E denuded. page 80 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Subfamily Cymoinae Alcock, 1898 Cymo melanodactylus Dana, 1852 # Cymo melanodactylus — Stimpson 1858a: 34; Stimpson 1907: 59; Shen 1940a: 226 [no new specimens]. Cymo memanodactylus — Gee 1926: 162 [list]. Taxonomy: Sakai (1939), Serène (1984), Holthuis (1993), Brösing et al. (2014). Type: Syntype ♂ RMNH D 42164 (Holthuis 1993; Fransen et al. 1997). Type locality: Java, Indonesia. Distribution: Western Indian Ocean to Central Pacific: from Madagascar, Seychelles, Mauritius, Red Sea, Bay of Bengal, to Southeast and East Asia to Japan, and Australia, Gilbert Islands, Fiji, Samoa, Tahiti and Tuamotu (Serène 1984). Habitat: Subtidal; associated with coral reefs (Dai and Yang 1991). Subfamily Etisinae Ortmann, 1893 Etisus anaglyptus H. Milne Edwards, 1834 # (Figs. 23, 38F) Etisus anaglyptus — present record. Taxonomy: Sakai (1939: as E. (Etioides)), Serène (1984). Type: Possibly in MNHN (Davie 2002). Type locality: “habite l'australasie”: in Australasia. Distribution: Western Indian Ocean to West Pacific: from Madagascar, Somalia, Red Sea, Persian/ Arabian Gulf, to Phuket (Thailand), Southeast and East Asia to Japan, and Australia (Serène 1984; Galil and Vannini 1990; Dai and Yang 1991; Ng and Davie 2002). Habitat: Intertidal and shallow subtidal; under stones (Dai and Yang 1991). Remarks: This species is a new record for the fauna of Hong Kong. Fig. 22. Pilodius miersi (Ward, 1936) (male, 7.7 × 5.4 mm, Bluff Island): A, carapace; B, maxilliped 3; C, right chela; D, right P5; E, pleon; F, left G1, mesial view; G, tip of left G1, mesial view. Structures A to D denuded. page 81 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Etisus laevimanus Randall, 1840 # Etisus laevimanus — Gordon 1931: 528; Shen 1940a: 225 [no new specimens]; Morton and Morton 1983: fig. 12.9(10). Taxonomy: Sakai (1939), Serène (1984). Type: Syntypes ANSP CA3070 (Spamer and Bogan 1993). Type locality: Hawaii. Distribution: Western Indian Ocean to Central Pacific: from South Africa, Madagascar, Red Sea, Persian/Arabian Gulf, Arabian Sea, of Bengal, Singapore, East Asia to Japan, and Indonesia, Australia, New Caledonia, Gilbert Islands, Vanuatu, Fiji, Samoa, Tuamotu and Hawaii (Serène 1984). Habitat: Intertidal and subtidal; locally common among crevices or under rocks fragments. Subfamily Euxanthinae Alcock, 1898 Medaeops granulosus (Haswell, 1882) # Medaeus granulosus — Gordon 1931: 543, text-figs. 19, 22A; Shen 1940a: 224. ? Medaeus noelensis — Thompson and Shin 1983: tab. 4 [not Medaeus noelensis Ward, 1934). Medaeops granulosus — Poon 2000: 12. Taxonomy: Gordon (1931), Forest and Guinot (1961), Guinot (1967a), Serène and Umali (1972), Mendoza et al. (2009). Type: Lectotype ♀ in AM P40834 (designated by Mendoza et al. 2009). Type locality: Port Denison, Western Australia. Distribution: Eastern Indian Ocean and West Pacific: Western to northeastern Australia, Singapore, the Philippines, South China, Taiwan, Japan (Dai and Yang 1991; Davie 2002; Mendoza et al. 2009). Fig. 23. Etisus anaglyptus H. Milne Edwards, 1834 (male, 41.0 × 27.1 mm, Shelter Island): A, carapace; B, major chela; C, right P5; D, pleon; E, left G1, mesial view; F, left G1, lateral view; G, tip of left G1, lateral view. page 82 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Habitat: Intertidal and subtidal; substrates near shore of rock or stone (Dai and Yang 1991). Remarks: The only local record of Medaeus noelensis Ward, 1934, reported in Thompson and Shin (1983), collected from benthic fauna from sludges of the Victoria Harbour, is doubtful and is probably Medaeops granulosus. Medaeus noelensis (now in the genus Danielea Ng & Clark, 2003) is found in intertidal habitats and among rubble in rock pools (Ng and Clark 2003; Mendoza et al. 2014). Paramedaeus simplex (A. Milne-Edwards, 1873) Paramedaeus simplex — Horikoshi and Takeda 1982: tab. 1. Taxonomy: Guinot (1967a), Serène and Umali (1972), Takeda (1976), Serène (1984), Ng and Clark (2002). Type: Type material in Musée Godeffroy No. 5817c (original text), at present whereabouts and status unknown. Type locality: Madagascar. Distribution: Western Indian Ocean to Central Pacific: Madagascar, Seychelles, Red Sea, to Indonesia, the Philippines, Ryukyus (Japan), New Caledonia, Guam, Hawaii (Ng and Clark 2002). Habitat: Intertidal and shallow subtidal, under rubble (Ng and Clark 2002). Subfamily Kraussiinae Ng, 1993 Palapedia integra (De Haan, 1835) Kraussia integra — Gordon 1931: 527; Shen 1940a: 223 [no new specimens]. Kraussia nitida — Morton and Harper 1995: 37 [not Kraussia nitida Stimpson, 1858]. Paralepida nitida — Morton and Harper 1997: tab. 4, pl. 1L [not K. nitida Stimpson, 1858]. Taxonomy: Sakai (1939: as Kraussia), Serène (1972: as Kraussia), Ng (1993), Ko and Takeda (1999). Type: Holotype ♀ RMNH D 42202 (Yamaguchi and Baba 1993; Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: Eastern Indian Ocean and West Pacific: Northwestern Australia, Indonesia, Gilbert Islands; South China, Taiwan, Japan (Buitendijk 1960; Serène 1972); records in Hawaii remained doubtful (Sakai 1976; Castro 2011). Habitat: Intertidal and subtidal; in sand or under rocks (Dai and Yang 1991). Remarks: Morphological differences between P. integra (De Haan, 1835) and P. nitida (Stimpson, 1858) include morphology of the carapace: that of P. integra being broader, frontal region visible as four obscure lobes, while the carapace of P. nitida is slightly longer than broad and the front has four distinct triangular lobes (cf. Sakai 1976: pl. 101(3). The local record is based on material collected around Cape d’Aguilar presented in Morton and Harper (1997: pl. 1L) which conforms to P. integra. Subfamily Liomerinae Sakai, 1976 Liomera caelata (Odhner, 1925) Liamera caelata — J Jiang and Zhou 1982: 681. Taxonomy: Odhner (1925: as Carpilodes), Takeda and Koyama (1974), Takeda (1976). Type: Holotype in NHM; paratypes in NHMD, NHM and UUZM (original text). Type locality: Macclesfield Bank, South China Sea. Distribution: Eastern Indian Ocean to West Pacific: Cocos (Keeling) Islands, South China Sea, Japan, Palau, Kai and Murray Islands (Odhner 1925; Davie 2002). Habitat: Subtidal; associated with shallow coral reefs (Sakai 1976). Remarks: The only record of this reef-associated species from China and adjacent seas was that of Odhner (1925) from Macclesfield Bank in South China Sea, and no material was further reported despite decades of sampling efforts (Dai and Yang 1991; Yang et al. 2008). The only local record from musselascidian / algal zone (low intertidal zone) along Tolo Harbour presented by J. Jiang and Zhou (1982) remains uncertain. We treat it as a doubtful record. Liomera laevis (A. Milne-Edwards, 1873) Liomera laevis — Horikoshi and Takeda 1982: tab. 1. Taxonomy: Odhner (1925: as Carpilodes), Sakai (1939: as Carpilodes), Takeda (1976), Chen and Lan (1978). Type: Syntype probably in MNHN (Davie 2002). Type locality: New Caledonia. Distribution: Eastern Indian Ocean to Central Pacific: Cocos (Keeling) Islands, Phuket (Thailand), Southeast to East Asia to Japan; off northwestern Australia, Timor Sea, New Caledonia, Fiji and Moorea Islands (Odhner 1925; Sakai 1976; Davie 2002; Ng and Davie 2002). Habitat: Subtidal; in coral reefs in shallow waters (Sakai 1976). page 83 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Liomera margaritata (A. Milne-Edwards, 1873) Liomera margaritata — Horikoshi and Takeda 1982: tab. 1. Taxonomy: Odhner (1925: as Carpilodes), Sakai (1939: as Carpilodes), Serène (1984), SH Lee and Ko (2011). Type: Syntype probably in MNHN (Davie 2002). Type locality: New Caledonia. Distribution: Western Indian Ocean to West Pacific: Madagascar, Somalia, Red Sea, India, Mergui Islands, Southeast and East Asia to Japan, New Guinea, Australia, New Caledonia and Samoa (Serène 1984; Dai and Yang 1991; Davie 2002). Habitat: Intertidal and shallow subtidal; in rocky or coral reefs (Davie 2002). Liomera venosa (H. Milne Edwards, 1834) # Liomera venosa — Horikoshi and Takeda 1982: tab. 1; Morton and Morton 1983: 273, fig. 12.9(14). Taxonomy: Odhner (1925: as Carpilodes), Sakai (1939: as Carpilodes), Takeda (1976), Serène (1984). Type: Holotype ♂ MP-B 2350 sec in MNHN (Serène 1984). Type locality: Unknown; probably Mauritius (Davie 2002). Distribution: Eastern Indian Ocean to Central Pacific: Mergui Archipelago, Phuket (Thailand), Singapore, Vietnam, South China, Taiwan, Japan, to Palau, Indonesia, Australia, New Caledonia, and Tahiti; record in western Indian Ocean remained doubtful (Odhner 1925; Serène 1984; Poupin 1996; Ng and Davie 2002). Habitat: Intertidal and subtidal; along rocky shores, under stones or in crevices of rocks, to depths of 30 m (Sakai 1976). Subfamily Xanthinae MacLeay, 1838 Demania reynaudii (H. Milne Edwards, 1834) # (Figs. 24, 39A) ? Xantho reynaudi — Morton and Morton 1983: 273, fig. 12.9(11). Demania reynaudii — present record. Taxonomy: Guinot (1979), Garth and Ng (1985), Ng and Yang (1989). Type: Holotype ♀ MP-B3027S in MNHN (Guinot 1979). Type locality: “mer des Indes”: Indian Ocean, precise locality unspecified. Distribution: Indian Ocean to Southeast and East Asia: Indian Ocean (type locality), Vietnam, the Philippines, South China, Taiwan and Japan (Ng and Yang 1989). Habitat: Subtidal; associated with coral or rocky reefs. Remarks: Xantho reynaudii H. Milne Edwards, 1834 and X. scaberrima Walker, 1887 had long considered synonymous since Odhner (1925). Various past records in the region, including works of Sakai (1939), Buitendijk (1950) and Takeda and Miyake (1969a), had adopted this synonymy (see Garth and Ng 1985), until Guinot (1969a 1979) separated them. Both species had been recorded in local literature. The local record of X. reynaudii by Morton and Morton (1983) is based on a sketch of the animal (fig. 12.9(11)), but it is too schematic to be precisely identified. However, we have recently examined material identified as D. reynaudii from off Clear Water Bay, hence confirming its presence in the territorial seas. A large male individual (76.6 × 52.5 mm) is herein illustrated. The present species had been reported to be poisonous to humans if ingested (Alcala et al. 1988; Tsai et al. 1997), and this is probably also true for D. scaberrima (see Garth and Alcala 1977). Demania scaberrima (Walker, 1887) # Lophoxanthus scaberrimus — Balss 1922b: 126; Gee 1926: 163 [list]; Shen 1940a: 224. Demania scaberrima — Blackmore and Rainbow 2000: app. 1; Wong et al. 2021: 58, fig. 96, pl. 18C. Taxonomy: Guinot (1979), Garth and Ng (1985), Ng and Yang (1989). Type: Syntype probably in NHM (original text; Johnson 1970). Type locality: Singapore. Distribution: Eastern Indian Ocean to South China Sea and East Asia: East India, Andaman Sea, Montebello Island (Australia), to South China Sea and East Asia to Japan (Garth and Ng 1985; Jones and Berry 2000). Habitat: Subtidal; near shore substrates, rocky or sandy; depths 15 to 35 m (Dai and Yang 1991). Remarks: Likely to be a toxic species; see Remarks under D. reynaudii above. Leptodius affinis (De Haan, 1835) # Chlorodius exaratus — Stimpson 1858a: 34 [part: see Remarks below] [not Chlorodius exaratus H. Milne Edwards, 1834]. Leptodius exaratus — Balss 1922b: 127; Shen 1940a: 224; BKK Chan et al. 2005: 1, fig. 2 [not C. exaratus H. Milne Edwards, 1834]. page 84 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Xantho exaratus — Gordon 1931: 528, 544, text-figs. 20, 22B [not C. exaratus H. Milne Edwards, 1834]. Leptodius affinis — S-k Lee et al. 2013: 192, figs. 3, 4E–I; Bravo et al. 2021: tab. 2. Taxonomy: Sakai (1939: as Xantho (Leptodius) exaratus), S-k Lee et al. (2013). Type: Lectotype ♀ RMNH D 44644 and paralectotypes in RMNH (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997; S-k Lee et al. 2013). Type locality: Japan, locality unspecified. Distribution: Eastern Indian Ocean to Central Pacific: Laccadive Sea, Phuket (Thailand), Southeast and East Asia to Japan, to Indonesia and Australia, New Caledonia, Palau, Mariana, Caroline, Marshall, Gilbert Islands, and Fiji, Samoa, to Tuamotu (S-k Lee et al. 2013). Habitat: Intertidal and shallow subtidal; along rocky or pebbly shore, under stones and crevices of rocks, or in coral reefs. Remarks: Leptodius exaratus (H. Milne Edwards, 1834) sensu stricto has been verified to be found only in the western Indian Ocean, Persian/Arabian Gulf and Red Sea, and records eastwards under this name belong to L. affinis instead (S-k Lee et al. 2013). After the publication of preliminary results on collections from the NPEE (1858a) as “Chlorodius exaratus”, Stimpson (1907) presented detailed diagnosis of nine varieties (a to i), among which two, namely h, latus and i, granulosus, were recorded from Hong Kong. According to S.-k. Lee et al. (2013), these represented, respectively, now L. sanguineus (H. Milne Edwards, 1834) L. philippinensis Ward, 1941, or Macromedaeus distinguendus (De Haan, 1835). These two records are listed as separated entries below. Leptodius davaoensis Ward, 1941 # (Fig. 39B, inset) Leptodius davaoensis — present record. Taxonomy: Ward (1941), Yuan et al. (2021). Type: Holotype ♂ AMNH-8347 (Ward 1941). Fig. 24. Demania reynaudii (H. Milne Edwards, 1834) (male, 72.6 × 52.5 mm, off Clear Water Bay): A, carapace; B, frontal region and maxilliped 3, ventral view; C, right chela; D, left P5; E, thoracic sternites and pleon. page 85 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Type locality: Davao, the Philippines. Distribution: Southeast and East Asia: from Micronesia (Kosrae Island), Lombok (Indonesia), to Malaysia and Singapore, the Philippines, and South China (Yuan et al. 2021). Habitat: Intertidal and subtidal: beneath or among rock fragments in sheltered shores. Remarks: Taxonomy of the present species remained poorly known. Locally, this species is not rare along semi-sheltered sandflats and shallow subtidal habitats adjacent to coral communities, and identification of relevant material has been confirmed by J.C.E. Mendoza (ZRC). This species has recently been reported from Hainan (Yuan et al. 2021), and the presence in Hong Kong is not unexpected. Images showing live coloration of overall habitus and external surface of chela, is provided herein. This species is a new record for the fauna of Hong Kong. Leptodius gracilis (Dana, 1852) # Chlorodius gracilis — Stimpson 1858a: 32; Stimpson 1907: 56. Leptodius gracilis — Gee 1926: 163 [list: “near Hong Kong”]; Shen 1940a: 225 [no new specimens]. Taxonomy: Sakai (1939: as Xantho (Leptodius)), Forest and Guinot (1961), Serène (1984). Type: Presumably lost. Type locality: Wake Island, Central Pacific. Distribution: Western Indian Ocean to Central Pacific: East Africa, Mauritius, Seychelles, Red Sea, to Phuket (Thailand), to South China Sea and Japan, and Palau, Indonesia, Marshall and Gilbert Islands, Samoa, Tuamotu and Hawaii (Buitendijk 1960; Serène 1984). Habitat: Intertidal and shallow subtidal; along rocky shores or coral reefs. Leptodius sanguineus (H. Milne Edwards, 1834) ? Chlorodius exaratus var. h, latus Stimpson, 1907: 55, pl. 6(9). ? Leptodius latus — Shen 1940a: 225 [no new specimens]. Taxonomy: Sakai (1939: as Xantho (Leptodius)), Serène (1984). Type: Syntypes probably in MNHN (Davie 2002). Type locality: Mauritius. Distribution: Western Indian Ocean to Central Pacific: from South Africa, Madagascar, Mauritius, Red Sea, Persian/Arabian Gulf, Indian Ocean, to Southeast and East Asia, towards Palau, Indonesia, Australia, New Caledonia, Fiji, Samoa, Tuamotu and Hawaii (Buitendijk 1960; Serène 1984). Habitat: Intertidal and subtidal; rocky shores under stones, in crevices of rocks or shallow waters on coral reefs (Dai and Yang 1991). Remarks: Following the comments by S-k Lee et al. (2013), we tentatively place the record of var. latus described in Stimpson (1907) under L. sanguineus (see Remarks under L. affinis above). Liagore rubromaculata (De Haan, 1835) # Liagore rubromaculata — Alcock 1898: 93 [part: Hong Kong material only]; Kemp 1923: 408 [part: Hong Kong material only]; Gee 1926: 163 [list]; Gordon 1931: 528; Shen 1940a: 223; Morton and Morton 1983: 273, fig. 12.9(12); Blackmore and Rainbow 2000: app. 1; Wong et al. 2021: 59, fig. 97, pl. 18D. Taxonomy: Sakai (1939), Guinot (1971), Ng and Chen (2004b). Type: Lectotype ♂ RMNH D 42221 and paralectotypes in RMNH (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: West Pacific: Queensland (Australia), Indonesia, Vietnam, South China, Taiwan, and Japan (Guinot 1971; Ng et al. 2001; Davie 2002). Habitat: Subtidal; in coral reefs or near rocky coast; depths 15 to 30 m (Dai and Yang 1991). Macromedaeus adelus Mendoza, 2021 # (Fig. 25A–D) Xantho distinguendus — Gordon 1931: 528, 543, text-figs. 21, 22C; Forest and Guinot 1961: 57, pl. 1(3). Macromedaeus adelus — present record. Taxonomy: Mendoza (2021). Type: Holotype ♂ ZRC 2017.0465 and paratypes in ZRC. Type locality: Punggol End, Singapore. Distribution: South China Sea: so far from Singapore (Mendoza 2021) and South China (present study). Habitat: Intertidal; middle to low tidal levels, under rock fragments. Remarks: In describing Macromedaeus adelus from Singapore and providing an overview of the genus, Mendoza (2021) revisited Buitendijk’s (1960) doubts about the identities of Xantho distinguendus reported by Gordon (1931), in comparison with then syntypes of C. (X.) distinguendus. Gordon (1931) commented on the morphological similarities in the specimens of Medaeops granulosus (as Medaeus), Leptodius affinis (as Xantho exaratus) and Macromedaeus distinguendus (as Xantho) collected from Hong Kong. The material of the latter species was described as “(g)ranulation of chelipeds almost obsolete”, “(u)pper margin of merus of walking-legs not crested” and “(carpus and propodus of walking-legs) hardly any trace of the granulation” page 86 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan (p. 544). The morphologies of the chelae and ambulatory legs described as such differ substantially with that of M. distinguendus, which are common along intertidal mudflats of Hong Kong (Fig. 25E–H). The second author recently reexamined material deposited at the NHM previously examined by Gordon, with the catalogue number NHM 1930.12.2.150, collected by Barney from Hong Kong. The lot comprises three males (two adults, one juvenile) and nine females (three ovigerous). A large male (10.0 × 6.2 mm) is illustrated herein. In comparison with M. distinguendus, which is common in Japan, Korea and North China (Fig. 25E; e.g., Shen 1932b; Sakai 1939 1976; Kim 1973; Yuan et al. 2022), both adult males of Gordon’s material have ambulatory legs with carpi and propodi not crested (Fig. 25D; vs. Fig. 25H), and frontal lobes medially separated by a deep V-shaped cleft (Fig. 25A; vs. Fig. 25E). The surface of the major chela in these specimens is externally granular on upper half and nearly smooth on lower portion (Fig. 25B), in contrast with M. distinguendus being entirely coarsely granular and dorsally rugose (Fig. 25F). The features observed in Gordon’s (1931) material confirm the presence of M. adelus in Hong Kong. Macromedaeus distinguendus, as Serène (1984) noted (p. 178), appears to be distributed in the Sino-Japanese area in temperate-subtropical East Fig. 25. Macromedaeus adelus Mendoza, 2021 (male, 10.0 × 6.2 mm, Hong Kong; in NHM 1930.12.2.150): A, overall habitus; B, major chela; C, minor chela; D, P4 and P5. Macromedaeus distinguendus (De Haan, 1835) (male, 27.4 × 17.4 mm, Tung Chung): E, overall habitus; F, major chela; G, minor chela; H, P4 and P5. page 87 of 155 Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Asia. Concerning the two recently described, and apparently tropical M. adelus and M. hainanensis Yuan, Jiang & Sha, 2022, both showing overlapping geographical reported thus far (from northern South China Sea towards the equatorial Southeast Asia), recent studies by the second author (PKLN) and J.C.E. Mendoza (ZRC) has concluded both names being synonyms, with M. adelus enjoying priority, that M. adelus published on 5 November 2021 while M. hainanensis was released on 18 January 2022. Our present comparisons agree with this. Nevertheless, the presence of both sub-tropical to temperate (M. distinguendus) and tropical (M. adelus) lineages in Hong Kong is not surprising. Macromedaeus distinguendus (De Haan, 1835) # (Fig. 25E–H) Chlorodius distinguendus — Stimpson 1858a: 34. ? Xantho distinguendus — Alcock 1898: 113 [part?]. ? Chlorodius exaratus var. i, granulosus Stimpson 1907: 56, pl. 6(3). ? Leptodius exaratus var. granulosus — Gee 1926: 163 [list]. ? Leptodius granulosus — Shen 1940a: 225 [no new specimens]. Xanthodius distinguendus — Shen 1940a: 224. Macromedaeus distinguendus — Blackmore and Rainbow 2000: app. 1; Poon 2000: 12. Taxonomy: Shen (1932b; as Xantho), Sakai (1939: as Xantho), Forest and Guinot (1961: as Xantho), Guinot (1967c). Type: Lectotype ♂ RMNH D 44657 and paralectotypes in RMNH (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: Probably only in East Asia (Serène 1984): so far confirmed from coasts of China from Guangdong and Shandong, Korea, and Kyushu, Japan (Mendoza 2021). Habitat: Intertidal; under stones or in crevices of rocks (Dai and Yang 1991). Remarks: Following indications by S.-k. Lee et al. (2013), records of Chlorodius exaratus var. granulosus described in Stimpson (1907) are tentatively placed under M. distinguendus. The resolution of the actual identities of the various forms recognized by Stimpson (1858b 1907) from East Asia will require more collections and comparisons of characters (see also Remarks under Leptodius affinis and M. adelus above). Subfamily Zosiminae Alcock, 1898 Atergatis dilatatus De Haan, 1835 Atergatis dilatatus — Shen 1940a: 224. Taxonomy: Buitendijk (1960), Serène (1984), Holthuis (1993). Type: Holotype RMNH D 44480 (Holthuis 1993; Fransen et al. 1997). Type locality: “ex Mari Chinensi originem ducunt”: Chinese seas. Distribution: Eastern Indian Ocean to West Pacific: Sri Lanka and Bay of Bengal, Southeast and East Asia to Japan, Queensland (Australia), New Caledonia and Samoa (Serène 1984; Davie 2002; Ng and Davie 2002; Richer de Forges and Ng 2006). Habitat: Shallow subtidal; associated with coral reef (Davie 2002). Atergatis floridus (Linnaeus, 1767) # Atergatis floridus — Morton and Morton 1983: 273, fig. 12.9(7). Taxonomy: Ng and Davie (2007). Type: Neotype ♂ ZRC 1999.0332 (designated by Ng and Davie 2007). Type locality: Pulau Seringat, Singapore (neotype). Distribution: West to Central Pacific: verified from Singapore, the Philippines, Taiwan, Guam, Indonesia, New Guinea and Australia (Ng and Davie 2007), and Hawaii (Castro 2011). Habitat: Intertidal and subtidal; associated with coral reefs in shallow waters (Dai and Yang 1991). Remarks: The present species was reported to be toxic with tetrodotoxin and paralytic shellfish toxin (Noguchi et al. 1983; Llewellyn and Endean 1991; Tsai et al. 1997). Atergatis integerrimus (Lamarck, 1818) # Atergatis integerrimus — Stimpson 1858a: 32; Stimpson 1907: 42; Gee 1926: 162 [list]; Gordon 1931: 528; Shen 1940a: 224. Etisus sp. — Morton and Ruxton 1997: 1 unnumb. fig. (23). Taxonomy: Sakai (1939), Buitendijk (1960), Serène (1984). Type: Status unknown (Davie 2002). Type locality: Unknown. Distribution: Western Indian Ocean to West Pacific: from East Africa, Mauritius, Persian/Arabian Gulf, India, Andaman and Mergui Islands, Phuket (Thailand), Southeast and East Asia to Japan, Australia, New Caledonia and Samoa (Serène 1984). Habitat: Subtidal; on rocky substrates, of 10 to 30 m deep (Dai and Yang 1991). Atergatis latissimus (H. Milne Edwards, 1834) ? Atergatis frontalis — Shen 1940a: 224. Taxonomy: Sakai (1939: as A. latissimus frontalis), page 88 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Tweedie (1947), Serène (1984). Type: Holotype ♀ MNHN 4125 (Serène 1984). Type locality: Australia (as Nouvelle-Hollande). Distribution: Western Indian Ocean to Central Pacific: from Madagascar, Mauritius, Christmas Island, to South China and Japan, to Sumatra and Marshall Islands (Serène 1984). Habitat: Shallow subtidal; coral reef-associated (Sakai 1976). Remarks: C.-J. Shen (1940a) listed four species of Atergatis for Hong Kong. Of these, species A. frontalis (De Haan, 1835) is now regarded as a junior synonym of A. latissimus (H. Milne Edwards, 1834) (Tweedie 1947; Serène 1984; Davie 2002; Ng et al. 2008), a decision we follow herein (though note reservations expressed by De Man 1926). The record of “A. frontalis” by Shen (1940a), however, has some problems. This author, in the same year, listed also this species in a checklist of South China (Shen 1940c), citing De Man’s (1879: 85) record from Amoy (= Xiamen), noting also that he “examined some specimens from Hongkong”. This record from Amoy, however, was later shown to be A. reticulatus De Haan, 1835 instead (De Man 1926). The unanswered questions are whether Shen (1940a) followed De Man’s (1879) original record of “A. frontalis” in listing the species and missed or misread his later correction of the identification (De Man 1926), and/or he actually had examined specimens of both species himself. As both species occur in the western Pacific, we retain both of Shen’s (1940a) records of A. reticulatus and A. frontalis (= A. latissimus) for the moment. Atergatis reticulatus (De Haan, 1835) Atergatis reticulatus — Gordon 1931: 528; Shen 1940a: 224. Taxonomy: Sakai (1939), Buitendijk (1960). Type: Lectotype ♀ in RMNH D 944, and paralectotypes in RMNH (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: East Asia: South China, Taiwan, Korea and Japan. Habitat: Intertidal and subtidal; in crevices of rocks, to depths of 20 m (Dai and Yang 1991). Remarks: See Remarks under A. latissimus above. Atergatopsis germaini A. Milne-Edwards, 1865 # (Figs. 26, 39C) Atergatopsis germaini — present record. Taxonomy: Buitendijk (1960), Sakai (1976). Type: Syntypes in MNHN and probably also RMNH (Buitendijk 1960; Guinot 1964). Type locality: Côn Sơn Island (as Poulo-Condore), Vietnam. Distribution: West Pacific: New Guinea, the Philippines, Vietnam, East Asia to Japan (Buitendijk 1960; Sakai 1976; S-k Lee et al. 2008). Habitat: Shallow subtidal; rocky reefs near coral communities. Remarks: This species had been reported to be toxic, containing human-lethal levels of paralytic shellfish poison by Tsai et al. (1996). This species is a new record for the fauna of Hong Kong. Lophozozymus pictor (Fabricius, 1798) # Lophozozymus epheliticus — Shen 1940a: 224. Taxonomy: Sakai (1976), Ng and Holthuis (1993), Ng and Chia (1997). Type: Lectotype ♂ ZMUC 109-1 (now NHMD) (designated by Ng and Chia 1997). Type locality: Unknown. Distribution: Eastern Indian Ocean to South China Sea and East Asia: Western Australia, Southeast and East Asia to Japan (Ng and Chia 1997). Habitat: Intertidal and subtidal; under rocks or in crevices of coral reefs, to depths of 30 m (Dai and Yang 1991). Remarks: The local record of L. epheliticus by Shen (1940a), which cited Miers (1880) and L. octodentatus in Alcock (1898), should be referred to as L. pictor (cf. Ng and Chia 1997). Like several members of this subfamily, the present species had been reported to be toxic, with tetrodotoxin and paralytic shellfish poison in its tissues (Chia et al. 1993; Tsai et al. 1995). Subsection Thoracotremata Guinot, 1977 Superfamily Cryptochiroidea Paul’son, 1875 Family Cryptochiridae Paul’son, 1875 Cryptochirus coralliodytes Heller, 1860 # Cryptochirus coralliodytes — Wong et al. 2023: 9, figs. 5A, 8. Taxonomy: Fize and Serène (1957: as Troglocarcinus (Favicola) rugosus), Takeda and Tamura (1981: as Favicola rugosa); Kropp (1988a). Type: Lectotype ♀ in NMW; paralectotype ♂ in MNHN (designated by Kropp 1988a). Type locality: Red Sea. Distribution: Western Indian Ocean to Central Pacific: from Red Sea to Phuket (Thailand), Vietnam (Fize and Serène 1957), South China, Taiwan and West page 89 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan (designated by Shahdadi and Schubart 2017). Type locality: Singapore. Distribution: South China Sea: Peninsular Malaysia, Singapore, Gulf of Thailand, Borneo and South China (Dai and Yang 1991). Habitat: Supratidal; under rocks beneath shades of coastal vegetation. Remarks: In describing subgenera Parasesarma and Perisesarma, De Man (1895) defined the former as having the lateral margin entire, while the latter has an epibranchial tooth after the extra-orbital tooth. Subsequently, however, this distinction between the two taxa was long considered unreliable (Shen 1932b; Tweedie 1940; von Hagen 1978), and both genera were shown to be non-monophyletic groups (Shahdadi and Schubart 2017). As such, Perisesarma has been restricted to its type species Pe. dusumieri H. Milne Edwards, 1853. The West African species were included in new genus, Guinearma; Pe. fasciatum transferred into another new genus, Fasciarma; and all others referred to Parasesarma (Shahdadi and Schubart 2017). Haberma tingkok Cannicci & Ng, 2017 # Haberma tingkok Cannicci & Ng, 2017: 68, figs. 1–5; Bravo et al. 2021: tab. 2. Taxonomy: Cannicci and Ng (2017). Type: Holotype ♂ ZRC 2016.620, and paratypes in ZRC, SWIMS and MZUF. Type locality: Ting Kok mangroves, Tai Po, Hong Kong. Distribution: South China: so far from Hong Kong only. Habitat: Intertidal; tree-climbing on mangroves Kandelia obovata Sheue, Liu & Yong, 2003 and Aegiceras corniculatum (L.) Blanco, 1837, at heights 1.5 to 1.8 m above substrate (Cannicci and Ng 2017). Nanosesarma minutum (De Man, 1887) # Sesarma (Sesarma) gordoni Shen, 1935: 27, text-fig. 7; Shen 1940a: 236. Sesarma minutum — Hill et al. 1975: 59; Morton 1976: 103. Nanosesarma minutum — Soh 1978: 10, pl. 3a; Hill et al. 1978: 84; J Jiang and Zhou 1982: 676; ZG Huang and Mak 1982: app.; Melville and Morton 1983: 18, 19 (1 un-numb. fig.); Morton and Morton 1983: 93, figs. 7.3(8), 9.4(8); Britton 1990: tab. 1; Ong Che and Morton 1992: 222; ZG Huang and Lin 1993: app.; Morton and Harper 1995: 54, fig. 8(23); Z Huang et al. 1999: tab. 1; VCS Lai et al. 2006: 61, 2 unnumb. figs.; Bravo et al. 2021: tab. 2. Nanosesarma gordoni — Ong Che and Morton 1992: tab. 1. Taxonomy: Shen (1935: as Sesarma (S.) gordoni), Sakai (1939: as S. (S.) gordoni), Serène and Soh (1970). Type: Syntype probably in ZMG (“Zoologischen Museums in Göttingen”; De Man 1888a): status unknown. The ZMG collections have been incorporated into the SMF. Type locality: Island of Edam near Jakarta, Indonesia. Distribution: Eastern Indian Ocean to South China Sea and East Asia: Phuket (Thailand), Singapore, Indonesia, China, Taiwan and Japan; exotic species in New Zealand (Ahyong and Wilkens 2011) and Hawaii (Davie 1998; Carlton and Eldredge 2009; Castro 2011), possibly through ballast waters (Brockerhoff and McLay 2011). Habitat: Intertidal; among fouling or oyster clumps. Remarks: The species Sesarma (Sesarma) gordoni has been considered to be a junior synonym of minutum by various authors (Serène and Soh 1970; Davie 1998; Naderloo and Türkay 2009). Records of S. minutum in the Indian Ocean have been shown to be N. sarii Naderloo and Türkay, 2009 (Persian/ Arabian Gulf to Pakistan) and N. jousseaumei (Nobili, 1906) (Madagascar, Tanzania to Persian/Arabian Gulf) (Holthuis 1977; Naderloo and Türkay 2009) instead. Nanosesarma pontianacense (De Man, 1895) # (Figs. 29, 40A) Nanosesarma pontianacense — Bravo et al. 2021: tab. 2. Taxonomy: Serène (1967: as N. tweediei), Dai and Song (1977). Type: No longer extant: holotype ♀ in Naturhistorischen Museums in Lübeck (original text), but original collections destroyed in WWII (Sabaj 2016). Type locality: Pontianak, Sarawak, Indonesia. Distribution: Eastern Indian Ocean to South China Sea: From Phuket (Thailand) to Singapore, Malaysia, Borneo, Vietnam, and South China (Serène 1967; Ng and Davie 2002). Habitat: Intertidal; among oyster clumps on rocks. Remarks: The present species was recently reported in a press release by the Nature Conservacy, based on specimens collected on oyster reefs at Lau Fau Shan, northwestern New Territories (online resource: https://www.tnc.org.hk/en-hk/what-we-do/hong-kongprojects/a-tiny-crab-in-hong-kong-is-showing-bigpromise-for-oyster-reefs/; accessed March 2020). We herein provide illustrations of specimens previously collected from Tung Chung. The second author has examined specimens from Hong Kong, and they are conspecific with material from Southeast Asia. The types are no longer extant and a neotype will be need to page 96 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan be designated as this peculiar species will to be referred to its own genus at a later date (see also Ng et al. 2008: 224). Neosarmatium indicum (A. Milne-Edwards, 1868) # Neosarmatium punctatum — Morton 1976: 105; Soh 1978: 10, pl. 3b; Morton and Morton 1983: 214, fig. 11.4(6) [not Metagrapsus punctatus A. Milne-Edwards, 1873]. Neosarmatium indicum — Davie 1994: 43, figs. 1E, F, 4, 17; Bravo et al. 2021: tab. 2. Neosarmatium smithi — WPW Kwok and Tang 2005: 3, fig. 6. Taxonomy: Davie (1994), Ng et al. (1997). Type: Lectotype ♂ MNHN-B10927 (designated by Davie 1994). Type locality: Sulawesi, Indonesia. Distribution: Southeast and East Asia: Indonesia, Singapore, Borneo (Malaysia), the Philippines to South China (Davie 2002; Schubart and Ng 2002). Habitat: Supratidal; under tidal and freshwater influences, in deep burrows on muddy substrates. Remarks: Material reported as N. punctatum in Soh (1978) was verified to represent N. indicum instead (Davie 1994). Orisarma dehaani (H. Milne Edwards, 1853) # Sesarma dehaani — Stimpson 1858b: 106; Stimpson 1907: 134; SY Lee 1993: 203; TH Lui et al. 2002: app. 2. Holometopus dehaani — Hill et al. 1975: 77; Hill et al. 1978: 108; Soh 1978: 10, pl. 2d; Morton 1979a: 121; Morton and Morton 1983: fig. 11.4(4). Sesarma (Holometopus) dehaani — SY Lee and Leung 1999: 61, pl. 2. Sesarma plicata — Tam and Wong 2000: 125, 2 unnumb. figs. Chiromantes dehaani — WPW Kwok and Tang 2005: 3, fig. 7; Komai and Ng 2013: 540. Orisarma dehaani — Bravo et al. 2021: tab. 2. Taxonomy: Shen (1932b: as Sesarma (Holometopus)), Komai and Ng (2013: as Chiromantes), Schubart and Ng (2020). Type: Lectotype RMNH D 157, and paralectotypes in RMNH, NHM, NHMD and MNHN (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997; Komai and Ng 2013). Type locality: Japan, locality unspecified. Distribution: East Asia: Coasts of China, Taiwan, Korea and Japan. Habitat: Supratidal; burrowing along muddy riverbanks or under shades of coastal vegetation. Remarks: Various authors have noted that Chiromantes Gistel, 1848 was polyphyletic (Ng and Fig. 29. Nanosesarma pontianacense (De Man, 1895) (male, 5.3 × 6.4, Tung Chung): A, carapace; B, right chela; C, right cheliped dactylus, dorsal view; D, left P5. All structures denuded. page 97 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Liu 1999; Ng et al. 2008; Komai and Ng 2013), with the taxonomic value of the form of lateral margin of carapace, i.e., presence or absence of tooth behind extra-orbital angle, as adopted by Serène and Soh (1970) in genera classification, being questioned (as with Perisesarma and Parasesarma, cf. Shahdadi and Schubart 2017). Chiromantes and Pseudosesarma were recently revised by Schubart and Ng (2020), and a number of East Asian species, including the locally present C. dehaani, P. patshuni, Sesarmops sinensis and S. intermedia, have been transferred to Orisarma. Orisarma intermedium (De Haan, 1835) Sesarma intermedia — Stimpson 1858b: 105; Heller 1865: 64; Stimpson 1907: 133; Morton 1979a: 118. Sesarma intermedium — Gee 1926: 164 [list]; Hill et al. 1975: 65; Hill et al. 1978: 92. Sesarma (Sesarma) intermedia — Gordon 1931: 528; Shen 1940a: 236. Sesarmops intermedium — Soh 1978: 11, pl. 4b. Orisarma intermedium — Bravo et al. 2021: tab. 2. Taxonomy: Sakai (1939: as Sesarma (Sesarma)), Schubart and Ng (2020). Type: Lectotype RMNH D 165, and probable paralectotype in RMNH (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: Eastern Indian Ocean to South China Sea and East Asia: Mergui Islands, Indonesia, to South China, Taiwan, Korea and Japan (De Man 1888b; Sakai 1976). Habitat: Supratidal; along banks of stream banks leading to the shore, rather tolerant to freshwater. Remarks: See Remarks under Orisarma dehaani above. Orisarma patshuni (Soh, 1978) # Pseudosesarma patshuni Soh, 1978: 14, figs. e, f, pls. 1c, f, 3f; Morton and Morton 1983: 214, fig. 11.4(7); WPW Kwok and Tang 2005: 3, fig. 16. Orisarma patshuni — Bravo et al. 2021: tab. 2. Taxonomy: Soh (1978: as Pseudosesarma), Schubart and Ng (2020). Type: Holotype ♂ BM 1976:108 in NHM, and paratypes probably in ZRC. Type locality: Shui Hau, Lantau Island, Hong Kong. Distribution: South China, probably near estuaries of Pearl River: so far reported from Macau and Hong Kong, and Dapeng District, Shenzhen (Huang and Mao 2021). Habitat: Supratidal; on among rock fragments along stream backs leading to the shore, appears rather tolerant to freshwater. Remarks: See Remarks under Orisarma dehaani above. Orisarma sinense (H. Milne Edwards, 1853) # Sesarma sinensis — Stimpson 1858b: 105; Stimpson 1907: 133; Gee 1926: 165 [list]; Morton 1979a: fig. 6.19; SY Lee 1993: tab. 1. Seasarma (Sesarma) sinensis — Shen 1931b: 196, text-fig. 13, pl. 14(2, 3). Sesarma (Sesarma) sinensis — Shen 1940a: 236. Sesarmops sinensis — Morton 1976: 105; Soh 1978: 11, pl. 4c; Morton 1979a: 121; Morton and Morton 1983: 158, fig. 11.11(12); Fong et al. 2005: 63, 3 unnumb. figs.; WPW Kwok and Tang 2005: 3, figs. 15, 18a, 20. Sesarma (Sesarmops) sinensis — SY Lee and Leung 1999: 61, pl. 1. Taxonomy: Shen (1931b: as Seasarma (Sesarma)), Schubart and Ng (2020). Type: Syntype ♂ in MNHN (see De Man 1887; Schubart and Ng 2020). Type locality: “Mers de Chine”: China, locality unspecified. Distribution: South China: from Guangdong to Zhejiang (Dai and Yang 1991). Habitat: Supratidal and intertidal; burrowing along muddy banks. Remarks: See Remarks under Chiromantes dehaani above. Parasesarma affine (De Haan, 1837) # Sesarma (Parasesarma) plicata — Balss 1922b: 155; Shen 1940a: 236 [not Ocypode plicatum Latreille, 1803]. Sesarma plicata — Gee 1926: 165 [list] [not Ocypode plicatum Latreille, 1803]. Parasesarma affinis — Soh 1978: 10, pl. 3c; Morton and Morton 1983: fig. 11.5; SY Lee 1993: tab. 1; WPW Kwok and Tang 2005: 3, figs. 12, 21. ? Parasesarma plicatum — Hill et al. 1975: 77; Morton 1976: 100; Hill et al. 1978: 108; Soh 1978: 11, pl. 3e; Morton and Morton 1983: fig. 10.3(1) [not Ocypode plicatum Latreille, 1803]. Sesarma (Parasesarma) affinis — SY Lee and Leung 1999: 62, pl. 4. Parasesarma pictum — So and Lui 2007: 35, 4 unnumb. figs. Parasesarma affine — Rahayu and Ng 2010: 8, figs. 5–7; Bravo et al. 2021: tab. 2. Taxonomy: Rahayu and Ng (2010). Type: Lectotype ♂ RMNH D 129, and paralectotypes in RMNH (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997; Rahayu and Ng 2010). Type locality: Japan, locality unspecified. Distribution: East Asia: coasts of China, Taiwan, Korea and Japan (Rahayu and Ng 2010). Habitat: Supratidal; under limited tidal influences, in burrows, sometimes under shades of coastal page 98 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan vegetation. Remarks: The taxonomy of the Parasesarma plicatum-group was revised by Rahayu and Ng (2010) and past records in East Asia under the name “P. plicatum” represent P. affine instead, with P. plicatum sensu stricto now known to be restricted to localities around Indian Ocean and Southeast Asia. Parasesarma continentale Shih, Hsu & Li, 2023 # Sesarma bidens — Heller 1865: 64 [part]; Bürger 1893: 628; Stimpson 1907: 134 [part; material from Hong Kong only]; Gee 1926: 164 [list]. [not Grapsus (Pachysoma) bidens De Haan, 1835]. Sesarma (Chiromantes) bidens — Gordon 1931: 528; Tam and Wong 2000: 115, 2 unnumb. figs. Sesarma (Perisesarma) bidens — Shen 1940a: 237. Chiromanthes bidens — Hill et al. 1975: 48; Morton 1976: 103; Hill et al. 1978: 92; Morton 1979a: 118; Soh 1978: 10, pl. 2a; Melville and Morton 1983: 11, 1 unnumb. fig. (12); Morton and Morton 1983: 220, fig. 11.11(8), pl. 19D; SY Lee 1993: tab. 1. Sesarma (Perisesarma) bidens — SY Lee and Leung 1999: 64, pl. 5. Perisesarma bidens — BKK Chan 2001: tab. 2; Poon 2002: 15, fig. 1; Fong et al. 2005: 61, 4 unnumb. figs.; WPW Kwok and Tang 2005: 3, fig. 13; So and Lui 2007: 34, 3 unnumb. figs; Bravo et al. 2021: tab. 2. Parasesarma continentale Shih, Hsu & Li, 2023: 11, figs. 4, 7F–H, 9C, D, 10D, 11D, 12B. Taxonomy: Shahdadi and Schubart (2017; on “Perisesarma”); Shih et al. (2023). Type: Holotype ♂ NCHUZOOL 17065, and paratypes in NCHUZOOL and ZRC. Type locality: Lieyu, Kinmen, Taiwan. Distribution: Vietnam and South China: shores of Vietnam, Hainan, Hong Kong, to Kinmen, Matzu and Fujian (Shih et al. 2023). Habitat: Supratidal and intertidal, among rock fragments, often under shades of mangroves. Remarks: See remarks under Fasciarma fasciatum above. The species “Perisesarma bidens” was recorded from a good wealth of literature from various localities along the West Pacific, including Hong Kong. This was recently shown to be a species complex comprise of at least five lineages, each warrant species-level status, with Taiwan situated at a point of overlap (Shahdadi et al. 2022). Based on molecular and morphological evidences, these lineages are now recognized as separate species: P. bidens (De Haan, 1835) sensu stricto from Korea and the Japanese Archipelago, as south as Iriomote; P. cricotum Rahayu and Davie, 2002 from Indonesia and Philippines; P. sanguimanus Li, Shih & Ng, 2019 and P. insulare Shih, Hsu & Li, 2023 from Taiwan to Philippines; P. chiahsiang Shih, Hsu & Li, 2023 from Taiwan, especially Penghu Islands; and P. continentale along shores of Vietnam to South China (Shih et al. 2023). These species show subtle morphological distinctions, as well as distinctive live colorations (Shih et al. 2023). So far only P. continentale is found in Hong Kong. Parasesarma maipoense (Soh, 1978) Chiromanthes maipoensis Hill et al. 1975: 77 [nomen nudum]. Chiromanthes maipoensis Morton 1976: 105 [nomen nudum]. Chiromanthes maipoensis Soh, 1978: 11, figs. a, b, pls. 1a, d, 2c; Hill et al. 1978: 108; Melville and Morton 1983: 11, 2 unnumb. figs. (13, 14); Morton and Morton 1983: fig. 11.11(3); SY Lee 1993: tab. 1. Sesarma (Perisesarma) maipoensis — SY Lee and Leung 1999: 65, pl. 6. Sesarma (Chiromanthes) maipoensis — Markham 1990: 560; Li 2003: 154 [no new specimens]. Sesarma maipoensis — Markham 1992: tab. 1 [list]. Perisesarma maipoense — Ng et al. 2010a: 239, figs. 1–4. Taxonomy: Soh (1978: as Chiromanthes), Ng et al. (2010a: as Perisesarma), Shahdadi and Schubart (2017). Type: Holotype ♂ BM 1976:106 in NHM, paratypes in ZRC (Ng et al. 2010a). Type locality: Mai Po marshes near Shum Chun River, Hong Kong. Distribution: Vietnam and South China: northern Vietnam, Macau and Hong Kong (Ng et al. 2010a). Habitat: Supratidal; on raised levees of drainage channels and drier bunds, or in the mangrove littoral zone (Ng et al. 2010a). Remarks: See remarks under Fasciarma fasciatum above. This species was long believed to a Hong Kong endemic until the species was also found in northern Vietnam (Ng et al. 2010a) and probably has a wider range than is known. Earlier use of this name in the literature was seen in faunal checklist compiled by Hill et al. (1975 1978), and an overview of mangrove crabs by Morton (1976). In both treatments, however, no diagnoses or descriptions were provided, hence considered as a nomen nudum. Parasesarma pictum (De Haan, 1835) # Sesarma picta — Bürger 1893: 626. Sesarma (Parasesarma) picta — Gordon 1931: 528; Shen 1940a: 237. Parasesarma pictum — Hill et al. 1975: 77; Morton 1976: 100; Hill et al. 1978: 92, 108; Soh 1978: 11, pl. 3d; Morton 1979a: 121, fig. 6.17; Morton and Morton 1983: 87, fig. 7.3(1), pl. 6; Britton 1990: tab. 1; WPW Kwok and Tang 2005: 3, fig. 11; VCS Lai et al. 2006: 60, 2 unnumb. figs.; Bravo et al. 2021: tab. 2. Sesarma pictum — Choi 1992: 901. Taxonomy: Shen (1932b: as Sesarma (Parasesarma)), Sakai (1939: as Sesarma (Parasesarma)), Maenosono and Naruse (2015). page 99 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Type: Lectotype ♂ RMNH D 45168, and paralectotypes in RMNH (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: East Asia: coasts of China, Korea, Taiwan, Ryukyus and Japan (Sakai 1976; Yang et al. 2008); the Indonesian record (from Celebes), presented in De Man (1887 1888b) was listed with doubt later in 1895 (De Man 1895: p. 185), and excluded by Widyastuti and Rahayu (2016). Habitat: Supratidal and intertidal; in along relatively sheltered rocky shores or in tide pools, and along shores of rock fragments. Parasesarma tripectinis Shen, 1940 # Parasesarma erythrodactylum — Poon and Chan 2001: 20. Parasesarma tripectinis — Rahayu and Ng 2005: 177; Bravo et al. 2021: tab. 2. Taxonomy: Shen (1940b: as Sesarma (Parasesarma)), Davie (1993: as P. acis), Rahayu and Ng (2005), Maenosono and Naruse (2015). Type: Whereabouts and status unknown. Type locality: Jimei, Fujian (as Tsimei, Fukien), China. Distribution: East Asia: coasts of China, Taiwan and Japan (Dai and Yang 1991; Davie 1993). Habitat: Supratidal to intertidal, usually in the upper zones; among coast vegetation and foliage, or under rocks. Parasesarma ungulatum (H. Milne Edwards, 1853) Parasesarma ungulatum — Rahayu and Ng 2010: 11, figs. 8–10; Bravo et al. 2021: tab. 2. Taxonomy: Rahayu and Ng (2010). Type: Lectotype ♂ MNHN BP 3694 (designated by Rahayu and Ng 2010). Type locality: Sulawesi, Indonesia. Distribution: Southeast and East Asia: Indonesia, Singapore, Malaysia, Thailand, to South China (Rahayu and Ng 2010). Habitat: Supratidal to intertidal; under mangroves. Sarmatium germaini (A. Milne-Edwards, 1869) # Sarmatium germanii — Hill et al. 1975: 77; Hill et al. 1978: 108. Sarmatium germaini — Soh 1978: 11, pl. 4a; Morton 1979a: 121; Morton and Morton 1983: 227, fig. 11.11(4); Bravo et al. 2021: tab. 2. Taxonomy: Serène and Soh (1971), Davie (1992c). Type: Lectotype ♀ MP-B3668 and paralectotypes in MNHN (designated by Davie 1992c). Type locality: Côn Sơn Island (as Poulo condore), southern Vietnam, South China Sea. Distribution: West Pacific: northern Australia, Malaysia, Singapore, the Philippines, Vietnam and South China (Davie 1992c). Habitat: Supratidal and intertidal; in burrows under shades of mangroves, some 10 m inland from the creek bank (Davie 1992c). Sarmatium striaticarpus Davie, 1992 Sarmatium striaticarpus — Bravo et al. 2021: tab. 2. Taxonomy: Davie (1992c). Type: Holotype ♂ ZRC 1970.1.23.14; and paratype ♀ in USNM. Type locality: Johore Straits. Distribution: South China Sea and East Asia: Singapore and Malaysia, the Philippines; Hong Kong (South China), to southern Okinawa (Davie 1992c; CGS Tan and Ng 1994; Bravo et al. 2021). Habitat: Intertidal; associated with mangrove swamps (Davie 1992c). Sinosesarma tangi (Rathbun, 1931) # Sesarma tangi — SY Lee 1993: tab. 1. Sesarma (Holometopus) tangi — SY Lee and Leung 1999: 62, pl. 3. Chiromantes tangi — WPW Kwok and Tang 2005: 3, fig. 10. Sinosesarma tangi — Ng et al. 2019: 4, figs. 1–5; Bravo et al. 2021: tab. 2. Taxonomy: Ng and Liu (1999: as Chiromantes), Schubart and Ng (2002: as Neosarmatium), Ng et al. (2019). Type: Holotype ♂ USNM 61875 (Ng and Liu 1999; Schubart and Ng 2002; Ng et al. 2019). Type locality: Guantao, near Fuzhou, Fujian (as Foochow, Fukien), China. Distribution: Vietnam and South China: northern Vietnam, Hong Kong and Fujian. Habitat: Supratidal to intertidal; burrowing near bases of trees in the interior of mangrove forests (SY Lee and Leung 1999; Ng et al. 2019). Family Varunidae H. Milne Edwards, 1853 Subfamily Asthenognathinae Stimpson, 1858 Asthenognathus hexagonum Rathbun, 1909 # Asthenognathus hexagonum — Wong et al. 2021: 60, fig. 98a–c, pl. 18E. Taxonomy: W Jiang et al. (2007), Yang and Tang page 100 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan (2008). Type: Holotype ♀ probably in NHMD (original text). Type locality: North of Koh Kong, Gulf of Thailand. Distribution: South China Sea: Gulf of Thailand, the Philippines and South China (Rathbun 1910; Serène and Soh 1976; Yang and Tang 2008). Habitat: Subtidal; muddy substrates; depths 8 to 26 m (Yang and Tang 2008). Asthenognathus inaequipes Stimpson, 1858 # Asthenognathus inaequipes — Ong Che and Morton 1991: tab. 1; Wong et al. 2021: 63, fig. 98d–f, pl. 18F. Taxonomy: De Man (1907), Sakai (1939), W Jiang et al. (2007). Type: Presumably lost. Type locality: “prop oras orientales insulæ “niphon”; lat. Bor. 38°”: seas off now Sendai, Japan. Distribution: East Asia: coasts of China, Korea, Japan (Sakai 1976; W Jiang et al. 2007; SH Lee et al. 2010). Habitat: Subtidal; substrates of mud or sandy mud; depths 7 to 50 m (W Jiang et al. 2007); also reported to be collected from dwelling of holothurian Protankyra bidentata (Woodward & Barrett, 1858) (SH Lee et al. 2010). Subfamily Cyclograpsinae H. Milne Edwards, 1853 Chasmagnathus convexus De Haan, 1835 # Chasmagnathus convexus — Balss 1922b: 154; Gee 1926: 164 [list]; Gordon 1931: 528; Shen 1940a: 235; Hill et al. 1978: errata; Morton 1979a: 121; SY Lee and Leung 1999: 66, pl. 7; Fong et al. 2005: 58, 4 unnumb. figs.; WPW Kwok and Tang 2005: 3, figs. 1, 17b, 18b, 19b, 22; K Sakai et al. 2006: 10, figs. 7–13; Bravo et al. 2021: tab. 2. Tiomanium mortoni Hill et al. 1975: 77 [nomen nudum]; Hill et al. 1978: 108. Chasmagnathus convexum — Melville and Morton 1983: 11, 2 unnumb. figs. (12, 14); Morton and Morton 1983: pl. 19E; SY Lee 1993: tab. 1. Taxonomy: Sakai (1939), K Sakai et al. (2006). Type: Lectotype ♂ RMNH D 213, and paralectotypes in RMNH (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: East Asia: South China, Taiwan, Korea and Japan (K Sakai et al. 2006). Habitat: Supratidal and intertidal; in burrows under shades of mangroves. Remarks: The names Tiomanium mortoni and T. shumchunensis appeared in a faunal checklist compiled by Hill et al. (1975 1978) but no diagnoses or comparisons were furnished for both taxa; they are therefore nomina nuda. In an errata sheet attached to the revised version of the checklist (Hill et al. 1978), these two names were indicated to instead represent Chasmagnathus convexus and Helice tientsinensis, respectively. Cyclograpsus incisus Shen, 1940 ? Cyclograpsus punctatus — Stimpson 1858b: 105; Stimpson 1907: 132; Gee 1926: 164 [list]; Shen 1940a: 236 [no new specimens] [not C. punctatus H. Milne Edwards, 1837]. Cyclograpsus incisus Shen 1940a: 236 [nomen nudum]. Cyclograpsus incisus Shen, 1940b: 259, figs. 10–16. Taxonomy: Shen (1940b). Type: Status unknown. Type locality: Aberdeen, Hong Kong. Distribution: South China: so far from Guangdong only. Habitat: Intertidal (Dai and Yang 1991). Remarks: Cyclograpsus punctatus H. Milne Edwards, 1837 has been recorded from an exceptionally wide geographic range, and the records almost certainly belong to complex of different species with C. punctatus sensu stricto found only in Chile and South Africa (Campbell and Griffin 1966; Griffin 1968b). Griffin (1968b) suggested that local records by Stimpson (1858b 1907) are C. incisus, a poorly understood species (see also Dai and Yang 1991). His recommendation is followed here. Cyclograpsus intermedius Ortmann, 1894 # Cyclograpsus intermedius — Morton and Morton 1983: 93, fig. 7.3(5). Taxonomy: Sakai (1939). Type: Syntypes in MZS 1294 (Komai 1999). Type locality: Amani-Oshima (Ryukyu, Japan), and Indian Ocean (locality unspecified). Distribution: Indo-West Pacific: Indian Ocean, and South China, Taiwan, Korea and Japan (Ortmann 1894). Habitat: Intertidal; beneath boulders and rock fragments. Helicana doerjesi K. Sakai, Türkay & Yang, 2006 Helicana doerjesi — Bravo et al. 2021: tab. 2. Taxonomy: K Sakai et al. (2006); Shih and Suzuki (2008). Type: Holotype ♂ SMF-8667; paratypes in ZIAS, BNHM, ZMH, USNM, SMF and ZIRAS. page 101 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Type locality: Mailiao, western Taiwan. Distribution: East Asia: Southern to Central China, and west coast of Taiwan (K Sakai et al. 2006; Shih and Suzuki 2008). Habitat: Intertidal; estuarine mudflats associated with mangroves. Helice latimera Parisi, 1918 # ? Helice tridens — Koelbel 1897: 711; Gee 1926: 164 [list]; Morton and Morton 1983: 227, fig. 11.11(7); SY Lee 1993: tab. 1 [not H. tridens De Haan, 1835]. Helice tridens pingi — Shen 1940a: 236. ? Tiomanium shumchunensis Hill et al. 1975: 77 [nomen nudum]; Hill et al. 1978: 108. ? Helice tientsinensis — Hill et al. 1978: errata [not H. tientsinensis Rathbun, 1931]. Helice latimera — SY Lee and Leung 1999: 66, pl. 8; K Sakai et al. 2006: 20, figs. 18, 22, 26, 30, 31; Bravo et al. 2021: tab. 2. Taxonomy: K Sakai et al. (2006). Type: Holotype ♀ MCSNM Nr. 1651 (K Sakai and Yatsuzuka 1980; Froglia and Grippa 1986). Type locality: China, locality unspecified. Distribution: Vietnam and South China: northern Vietnam and South China. Habitat: Intertidal; under beds of common reed Phragmites australis (SY Lee and Leung 1999). Remarks: On the name Tiomanium shumchunensis, see Remarks under Chasmagnathus convexus above. The record of “H. tientsinensis” is likely to be H. latimera instead; with K. Sakai et al. (2006) and Shih and Suzuki (2008) showing that the former species is distributed in North China and Korea instead. Helice sp. Helice sp. — SY Lee and Leung 1999: 67, pl. 9. Remarks: Based on material collected from the Mai Po Marshes, S.Y. Lee and Leung (1999) reported two species of Helice: H. latimera Parisi, 1918, and another form as Helice sp. The latter form, as commented by the late Prof. A.-Y. Dai, being “still obscure and may represent a new species”. Unfortunately, their treatment on Hong Kong sesarmid crabs (cited as “Lee, Dai and Chen, in press”) has never been published. This material is now in the collection of the Queensland Museum and will be re-examined and studied at a later date (S.Y. Lee, Peter Davie, pers. comm.). We retain this species record for the time being. Metaplax elegans De Man, 1888 Metaplax elegans — Gordon 1931: 528; Shen 1940a: 236; Davie 1992a: 352, pl. 2B; SY Lee and Leung 1999: 68, pl. 11; WPW Kwok and Tang 2005: 3, fig. 4. Taxonomy: De Man (1888b), Davie (1992a), Shih et al. (2019). Type: Syntypes RMNH D 13 (Fransen et al. 1997), NHM 1886.52 (NHM Data Portal), and probably also in ZSI (as Indian Museum, Calcutta in original text). Type locality: Mergui Archipelago, Myanmar. Distribution: Eastern Indian Ocean and South China Sea: East India, Mergui Archipelago, Phuket (Thailand), Malaysia, to South China and Taiwan (Shih et al. 2019). Habitat: Intertidal; on mudflats under mangroves. Metaplax longipes Stimpson, 1858 # (Fig. 30) Metaplax longipes Stimpson, 1858b: 97; Koelbel 1897: 711, pl. 1(1– 4); Stimpson 1907: 99; Gee 1926: 164 [list]; Gordon 1931: 528; Shen 1940a: 236; Shih et al. 2019: 9, figs. 2D–F, 4, 7E–H; Bravo et al. 2021: tab. 2. Macrophthalmus boteltobagoe — Bones 1982: 691, pls. 1, 2c, d; Morton and Morton 1983: 156 [not Ma. boteltobgoe Sakai, 1939]. Metaplax takahasii — Davie 1992a: 352, pl. 2A; SY Lee and Leung 1999: 69. ? Metaplax longipes — SY Lee 1993: tab. 1; SY Lee and Leung 1999: 68, pl. 10; WPW Kwok and Tang 2005: 3, figs. 5, 19a; So and Lui 2007: 36. Taxonomy: Stimpson (1907), Shih et al. (2019). Type: Presumably lost. Type locality: Hong Kong. Distribution: South China Sea and East Asia: northern Vietnam, South China, and west coast of Taiwan (Shih et al. 2019). Habitat: Intertidal; on mudflat along fringes of mangroves, often associated with estuarine conditions. Remarks: See remarks under Metaplax tredecim below. Local records of “Macrophthalmus boteltobagoe” require some discussion. This species was reported by Bones (1982) from Tong Fuk Miu Wan, and Morton and Morton (1983) from Shui (as Shiu) Hau, both sites on of sheltered sandflats on Lantau Island. The reported habitat was very different from earlier reports of Ma. boteltobagoe sensu stricto: dwelling in burrows on rocky shores of limestone substrates, from various sites in the Ryukyus, Japan (Kosuge 1991; Kosuge and Davie 2001), and possibly also at Lanyu Island (as Kôtôsyo), Taiwan, where the species was described (Sakai 1939). We agree with Shih et al. (2015) that on the basis of the reported habitat, this record is not true Ma. boteltobagoe. The illustrations of Bones’ (1982) record (pls. 1, 2c, d), in any case, substantially differ from the figures of Ma. boteltobagoe Sakai, 1939, and Ma. page 102 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan holthuisi Serène, 1973(b). Compared to figures of both species in Kosuge and Davie (2001): (1) the local specimen measured approximately 1.9 × 1.4 cm, which is substantially larger than both Macrophthalmus species (cw. generally less than 1 cm: see Kosuge and Davie 2001); (2) the extraorbital tooth in the local specimen appears truncated to squarish (Bones 1982: pl. 2c, d; reproduced as Fig. 26A, B), whereas in both Macrophthalmus species, it is triangular, rather protruding, and slightly to distinctly acute; and (3) in the local specimen, it is devoid of any traces of fine setae or tomentum on the inner surfaces of the fingers of the chela (Bones 1982: fig. 2d; reproduced as Fig. 26B). As such, this record is neither Ma. boteltobagoe nor Ma. holthuisi. Bones’ (1982) species closely resembles Metaplax in having the cheliped and fringes of the carapace nearly naked and the ambulatory legs are markedly more slender and elongated. The anterolateral lobes of Bones’ (1982) taxon, being truncate, quadrate and not clearly protruding, actually agree with of Me. longipes, a common species along the shores of Lantau Island. We provide images of an adult male specimen of Me. longipes, size 18.2 × 13.5 mm from Tung Chung, for comparison in figure 30C, D. As such, we are confident Bones’ (1982) “Macrophthalmus boteltobagoe” is Me. longipes instead. Metaplax tredecim Tweedie, 1950 # Metaplax tredecim — Shih et al. 2019: 16, figs. 2J–L, 6, 7M–P; Bravo et al. 2021: tab. 2. Taxonomy: Tweedie (1950), Shih et al. (2019). Type: Holotype ♂ ZRC 1971.10.13.1 (Davie and Nguyen 2003). Fig. 30. Metaplax longipes Stimpson, 1858. A, B: reproduction of illustrations as “Macrophthalmus boteltobagoe” by Bones (1982: pls. 2c, d), with permission from the Hong Kong University Press; C, D: male, 18.2 × 13.5 mm, Tung Chung). page 103 of 155 Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Type locality: Labuan, Borneo. Distribution: Southeast and East Asia: northern Borneo, Vietnam, to South China (Shih et al. 2019). Habitat: Intertidal; on mudflat along fringes of mangroves, often on substrates of coarse sands. Remarks: Based on published records, three Metaplax species have been reported locally: M. elegans, M. longipes, and M. takahasii (Davie 1992a). Recently, Shih et al. (2019) showed M. takahasii Sakai, 1939 is a junior synonym of M. longipes Stimpson, 1858, and had been incorrectly represented as “M. takahasii” in Davie (1992a). The third species present was identified as M. tredecim Tweedie, 1950. The presence in Hong Kong (South China) may mark the northern limit of this essentially Southeast Asian species. Subfamily Gaeticinae Davie & N.K. Ng, 2007 Gaetice depressus (De Haan, 1833) # Platygrapsus depressus — Stimpson 1858b: 104; Stimpson 1907: 128. Platynotus depressus — Heller 1865: 60. Gaetice depressus — Balss 1922b: 150; Gordon 1931: 528; Shen 1940a: 238; J Jiang and Zhou 1982: 681; Markham 1992: tab. 1 [list]; Morton and Harper 1995: 43, fig. 6(3); Poon and Chan 2001: 21; VCS Lai et al. 2006: 59, 3 unnumb. figs.; Davie and NK Ng 2007: 262, figs. 4–6; Bravo et al. 2021: tab. 2. Goetice depressus — Gee 1926: 164 [list]; Markham 1982: 361; Li 2003: 153 [no new specimens]. Hemigrapsus goetica — Hill et al. 1975: 59. Goetica depressa — Hill et al. 1978: 84. Gaetice depressa — Morton and Morton 1983: 93, figs. 7.3(6), 7.12(10); BKK Chan 2001: tab. 2. Taxonomy: Sakai (1939), Davie and NK Ng (2007). Type: Lectotype RMNH D 1226, and paralectotypes in RMNH and MNHN (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: East Asia: coasts of China, Taiwan, Korea and Japan (Sakai 1976). Habitat: Intertidal; under rock fragments. Gaetice ungulatus Sakai, 1939 Gaetice ungulatus — Davie and NK Ng 2007: 264. Taxonomy: Sakai (1939). Type: Whereabouts and status unknown. Type locality: Okinawa, Japan. Distribution: East Asia: coasts of China, Taiwan and Japan (Davie and NK Ng 2007). Habitat: Intertidal; under small rocks, substrates of coral-sand, close to river mouths (Davie and NK Ng 2007). Sestrostoma toriumii (Takeda, 1974) # Acmaeopleura toriumii — Davie 1992a: 350. Sestrostoma toriumii — Davie and NK Ng 2007: 267. Taxonomy: Itani et al. (2002: as Acmaeopleura), Davie and NK Ng (2007), S Lee et al. (2013). Type: Holotype ♂ NSMT-Cr 971. Type locality: Onagawa Bay, Miyagi, Japan. Distribution: East Asia: South China, Korea and Japan. Habitat: Intertidal; substrates of coarse gritty sand, in burrows of mud shrimp Upogebia major (De Haan, 1841) or inn-keeper worm Ochetostoma erythrogrammon Leuckart and Ruppell, 1828 (Davie and NK Ng 2007). Subfamily Varuninae H. Milne Edwards, 1853 Eriocheir hepuensis Dai, 1991 # Eriocheir hepuensis — Wong et al. 2021: 64, fig. 99, pl. 19A. Taxonomy: Guo et al. (1997), Naser et al. (2012). Type: Holotype ♂ AS GX899024A in IZCAS (Guo et al. 1997). Type locality: Hepu, Guangxi, China. Distribution: South China from Guangxi to Fujian (Guo et al. 1997) and northern Vietnam (Clark et al. 2023); exotic species in Persian/Arabian Gulf (Naser et al. 2012; Naderloo 2014 2017). Habitat: Freshwater to estuarine shallow seas, catadromous. Remarks: Two female specimens morphologically identifiable as E. hepuensis Dai, 1991 were collected from the local western estuarine waters (Wong et al. 2021). See also Remarks under E. japonica below. Eriocheir japonica (De Haan, 1835) # Eriocheir japonicus — Gordon 1931: 528; Shen 1940a: 237; Dudgeon 1999: 178, fig. 4.21A; Dudgeon 2003: 60, 3 unnumb. figs. ? Eriocheir sinensis — Hill et al. 1975: 48; Hill et al. 1978: 67; Morton 1979a: 121; Morton 1979b: tab. 7.1, fig. 7.5b; Hill and Phillipps 1981: 146, pl. 49f; Morton and Morton 1983: 214, fig. 11.4(2). Eriocheir japonica — T-Y Chan et al. 1995: 302, figs. 1A, 2C, 3C; Guo et al. 1997: 460, figs. 5, 6c, 6f, 7c; Yam et al. 2002: 18; Wong et al. 2021: 65, fig. 100, pl. 19B. Taxonomy: T-Y Chan et al. (1995), Guo et al. (1997), Naser et al. (2012), Ng et al. (2017a). Type: Lectotype ♂ MNHN 3382, and page 104 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan paralectotypes in RMNH, NHM, NHMD and MNHN (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997; Guo et al. 1997). Type locality: Japan, locality unspecified. Distribution: East Asia: South China, Taiwan, Korea and Japan. Habitat: Freshwater to estuarine shallow seas, catadromous; often submerged, on substrates of rock fragments, covered with foliage or aquatic vegetation. Remarks: The taxonomic understanding of Eriocheir species from South China shows some disagreement between morphological and molecular identification approaches. The present taxon from Hong Kong, which is not uncommon in local fresh water, intertidal and shallow marine habitats, is morphologically clearly identifiable as E. japonica (De Haan, 1835) (cf. Guo et al. 1997). The available genetic data, however, suggests individuals from South China (including Hong Kong and Macau) form a clade associated with E. hepuensis Dai, 1991 (cf. Naser et al. 2012). This matter was recently elaborated in detail by Ng et al. (2017a), and we report the local material as E. japonica, separate from the morphologically distinct E. hepuensis treated above. Hemigrapsus penicillatus (De Haan, 1835) # (Fig. 40C, D, inset) ? Heterograpsus penicillatus — Stimpson 1858b: 104 [part: Hong Kong material only]; Stimpson 1907: 126 [part: Hong Kong material only]. ? Brachynotus penicillatus — Koelbel 1897: 712, pl. 1(5, 6); Gordon 1931: 552. ? Hemigrapsus penicillatus — Gee 1926: 164; Shen 1940a: 237; Morton and Morton 1983: 246, fig. 11.18(13); VCS Lai et al. 2006: 57, 3 unnumb. figs. Hemigrapsus penicillatus — Bravo et al. 2021: tab. 2; present record. Taxonomy: Asakura and Watanabe (2005), Mingkid et al. (2006). Type: Lectotype ♂ RMNH D 200, and paralectotypes in RMNH and MNHN (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997); the type lots contain individual(s) of H. takanoi (Asakura and Watanabe 2005). Type locality: Japan, locality unspecified. Distribution: East Asia: verified from various localities in Taiwan, Japan, Korea and Pacific coast of Russia (Asakura and Watanabe 2005; S Lee et al. 2013; Marin 2013; Ng et al. 2017a); possible distribution in China remains to be verified; and doubtfully from Hawaii (Castro 2011). Habitat: Intertidal; under boulders on mudflats, estuaries, lagoons and sheltered beaches (Asakura and Watanabe 2005). Remarks: Past records under the name Hemigrapsus penicillatus (De Haan, 1835) were demonstrated by Asakura and Yamaguchi (2005) to consist of H. penicillatus sensu stricto and the morphologically similar H. takanoi Asakura & Yamaguchi, 2005, both species originally described from Japan. These two species differ more consistently, in larger males, by the size of setal patch in relation to the chela, and configuration of pigmentation on external maxillipeds, pterygostomian region, and, in some cases, the anterior thoracic sternites and pleon (see also Mingkid et al. 2006; Marin 2013). Using these criteria to examine the local material, we also recognize two subtly distinct forms: from eastern sites (e.g., in vicinity of Tolo Channel) where the setal patch on the male chela is relatively smaller, being approximately half of the chela height (Fig. 40D inset), the chela is externally covered by sparse dark dots with those on the third maxillipeds and pterygostomian regions slightly larger (Fig. 40D); whereas material from western sites (e.g., around Lantau Island), have a comparatively larger setal patch on the male chela and the external surface is grayish but not spotted (Fig. 40F inset), with the third maxillipeds and pterygostomian region covered with relatively smaller dark dots (Fig. 40F). This distinctness of both forms is supported by preliminary molecular results (HTS, unpublished data). These two forms are tentatively identified as H. penicillatus (eastern: Fig. 40C, D, inset) and H. takanoi (western: Fig. 40E, F, inset). They, however, differ slightly from their respective Japanese and Northeast Asian counterparts. For H. penicillatus, the local form had never been observed to be decorated with dark round spots as large, and the dark spots do not extend to the anterior thoracic sternites and pleon, as shown by Asakura and Yamaguchi (2005: figs. 8–10) and Marin (2013: fig. 2). As for H. takanoi, the local form has a relatively inflated chela palm with the external surface unspotted (vs. less inflated with with numerous scattered small dark dots; Asakura and Yamaguchi 2005: fig. 3; Marin 2013: fig. 1). As such, more morphological and molecular investigations are needed to be done on these two taxa in Hong Kong. Hemigrapsus takanoi Asakura & Watanabe, 2005 # (Fig. 40E, F, inset) Hemigrapsus takanoi — present record. Taxonomy: Asakura and Watanabe (2005), Mingkid et al. (2006), S Lee et al. (2013), Ng et al. (2017a). Type: Holotype ♂ CBM-ZC 8039, and paratypes in CBM. Type locality: Estuary of Okamoto-gawa, Tomiura, page 105 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan regarded as junior synonym of S. globosa De Haan, 1835 (since Koelbel 1897). Shen’s (1935) local record of S. tuberculata, provided with rather precise illustrations (and his comparison with Japanese material of S. globosa), matches with material of S. intermedia we collected from Hong Kong, western Taiwan and Singapore (the latter proximal to type locality, Johore; Wong et al. 2010a). Tmethypocoelis ceratophora (Koelbel, 1897) # Dioxippe ceratophora Koelbel, 1897: 714, pl. 1(8–12). Tympanomerus ceratophora — Gordon 1931: 528. Tmethypocoelis ceratophora — Shen 1935: 35, text-figs. 11–13; Shen 1940a: 233; Hill et al. 1975: 71; Hill et al. 1978: 92, 100; Morton 1979a: 121; Morton and Morton 1983: 185, fig. 10.3(2); Choi 1992: 901; Shea 1992: tab. 2; Davie and Kosuge 1995: 210, figs. 1A, C, E, 2A, B; BKK Chan 2001: tab. 2, fig. 1c; BKK Chan and Caley 2003: 90, 3 figs.; Fong et al. 2005: 55, 3 unnumb. figs.; Bravo et al. 2021: tab. 2. Taxonomy: Shen (1935), Davie and Kosuge (1995), Shih et al. (2015). Type: Whereabouts and status unknown. Type locality: Hong Kong. Distribution: East Asia: South China and Taiwan (Shih et al. 2015). Habitat: Intertidal; sandflats or under shades of mangroves, substrates of coarse or gritty sand. Subfamily Sheniinae Ng, Clark & Cuesta, 2010 Shenius anomalus (Shen, 1935) Camptandrium anomalum Shen, 1935: 31, text-figs. 8B, 9; Shen 1940a: 234. Shenius anomalus — Ng et al. 2010b: 1534, figs. 1–10. Taxonomy: Serène and Umali (1972), Ng et al. (2010b). Type: Holotype ♂ CAS CB-02060a, and paratype ♀ in IZCAS (Ng et al. 2010b). Type locality: Tai Po, Hong Kong. Distribution: South China Sea: Singapore, Peninsular Malaysia, to South China (Ng et al. 2010b). Habitat: Intertidal; exclusively in mangroves, under planks, wood or rafts on sandy mud (Ng et al. 2010b). Family Macrophthalmidae Dana, 1851 Subfamily Ilyograpsinae Števčić, 2005 Ilyograpsus paludicola (Rathbun, 1909) Ilyograpsus paludicola — Bravo et al. 2021: tab. 2. Taxonomy: Komai and Wada (2008). Type: Holotype ♀ ZMUC 7692 (Komai and Wada 2008). Type locality: Laem Ngop (as Lem Ngob), Gulf of Thailand. Distribution: West Pacific: Labuan (Malaysia), Singapore, Indonesia, Vietnam, to Hong Kong (South China), to northern Australia and New Caledonia (Komai and Wada 2008; Bravo et al. 2021). Habitat: Intertidal; associated with mangroves, on tidal flats and in estuarine habitats. Subfamily Macrophthalminae Dana, 1851 Remarks: The subgeneric classification of the genus Macrophthalmus Desmarest, 1823, as presented in Barnes (1967), Komai et al. (1995) and Ng et al. (2008) has been challenged by molecular phylogenetic data (Horii et al. 2001; Kitaura et al. 2006), and is still in need of revision. We follow Shih et al. (2015) in omitting subgeneric indications for the time being. For the local record of M. boteltobagoe presented by Bones (1982), see Remarks under Metaplax longipes above. Macrophthalmus abbreviatus Manning & Holthuis, 1981 # Macrophthalmus dilatatus — Shen 1940a: 235; Morton 1979a: 121. Macrophthalmus abbreviatus — Davie 1992a: 348 [key]; SY Lee and Leung 1999: 73. Taxonomy: Tesch (1915: as M. dilatatus), Shen (1932b: as M. dilatatus), Sakai (1939: as M. dilatatus), Shih et al. (2015). Type: Lectotype ♂ RMNH D 299, and paralectotype in RMNH and MNHN (designated by Yamaguchi and Baba 1993; see also Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: Eastern Indian Ocean to South China Sea and East Asia: Indonesia, coasts of China, Taiwan, Korea and Japan. Habitat: Intertidal; waterlogged substrates on sandflats or mudflats. Macrophthalmus banzai Wada & K. Sakai, 1989 Macrophthalmus banzai Wada & K Sakai, 1989: 133, figs. 1, 2; Davie 1992a: 348 [key]; SY Lee and Leung 1999: 73. Taxonomy: Wada and K Sakai (1989), Shih et al. (2015). Type: Holotype ♂ SMF 17923, and paratypes in page 112 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan SMF, SMBL, USNM, MNHN, RMNH and AM. Type locality: Samusaura, Shirahama, Wakayama, Japan. Distribution: East Asia: coasts of China, Taiwan, Korea and Japan. Habitat: Intertidal; waterlogged substrates on mudflats. Macrophthalmus convexus Stimpson, 1858 # Macrophthalmus convexus — Gordon 1931: 528; Shen 1940a: 235; Barnes 1970: 222; Morton 1976: 103; Bones 1982: 689, pl. 2a, b; Melville and Morton 1983: 11, 1 unnumb. fig. (12); Morton and Morton 1983: 156; Davie 1992a: 348 [key]; SY Lee 1993: tab. 1; Poupin 1997: fig. 3E; SY Lee and Leung 1999: 73; Bravo et al. 2021: tab. 2. Macrophthalmus spp. — Fong et al. 2005: 52, 3 unnumb. figs. Taxonomy: Komai et al. (1995), Davie (2012), Shih et al. (2015). Type: Probably lost. Type locality: Ryukyu Islands (as Loo Choo Isles), Japan. Distribution: Western Indian Ocean to Central Pacific: from Mauritius, India, Myanmar, Thailand, Peninsular Malaysia, South China Sea and East Asia, to Indonesia, Australia, New Caledonia, Fiji, French Polynesia and Hawaii (Shih et al. 2015). Habitat: Intertidal; in shallow burrows on sandymuddy substrates. Macrophthalmus definitus Adams & White, 1849 Macrophthalmus teschi — Gordon 1931: 549. Macrophthalmus definitus — Shen 1940a: 234; Bones 1982: 691: pl. 3a, b; Morton and Morton 1983: 157; Shin 1990: tab. 2; Davie 1992a: 348 [key]; SY Lee and Leung 1999: 73; Bravo et al. 2021: tab. 2. Macrophthalmus (Mareotis) definitus — Barnes 1970: 232. Taxonomy: Barnes (1967 1970), Komai et al. (1995), Shih et al. (2015). Type: Holotype NHM 43.6 (Barnes 1970). Type locality: The Philippines. Distribution: Eastern Indian Ocean to West Pacific: Phuket (Thailand), Peninsular Malaysia, the Philippines, South China, Taiwan, Ryukyus, and Indonesia to the Solomon Islands (Shih et al. 2015). Habitat: Intertidal; in burrows on very soft substrates on mudflats near mangrove swamps (Komai et al. 1995). Remarks: Based on unpublished notes by C.-J. Shen deposited at the British Museum, the local record of “M. teschi” in Gordon (1931) represents the present species instead (Barnes 1970). Macrophthalmus dentatus Stimpson, 1858 Macrophthalmus dentatus Stimpson, 1858b: 97; Stimpson 1907: 96, pl. 15(1); Gee 1926: 165 [list]; Shen 1940a: 235 [no new specimens]; Davie 1992a: 348 [key]. Taxonomy: Barnes (1971), Davie (2012). Type: Presumably lost. Type locality: Hong Kong. Distribution: West Pacific: Queensland (Australia), Timor, Gulf of Thailand and South China (Davie 2012). Habitat: Subtidal; coastal substrates of muddy sand; depths 8 to 20 m (Davie 2002). Macrophthalmus depressus Rüppell, 1830 Macrophthalmus depressus — Shen 1940a: 235. Taxonomy: Crosnier (1965: M. depressus sensu stricto), Barnes (1970), Naderloo et al. (2011). Type: Syntypes RMNH D 42227 (see Fransen et al. 1997). Type locality: Northern Red Sea. Distribution: Western Indian Ocean to Arabian Sea: South and East Africa, Madagascar, Red Sea, Persian/Arabian Gulf, Pakistan and western India (Naderloo et al. 2011). Habitat: Intertidal; on muddy and sandy-muddy substrates. Remarks: The local record of M. depressus Rüppell, 1830 was from Shen (1940a), who cited Alcock (1900b) whose material was from Mergui and Aden. Specimens from Mergui were later separated and described as M. teschi by Kemp (1919b), whereas the species M. depressus Rüppell, 1830 sensu stricto was subsequently restricted to the Indian Ocean and Red Sea (Stephensen 1946; Holthuis 1958; Crosnier 1965; Barnes 1970; Naderloo et al. 2011). The record of M. depressus from Hong Kong thus appears rather unlikely (Davie 1992b) but is retained here for the moment with doubt. Macrophthalmus erato De Man, 1888 # Macrophthalmus erato — Koelbel 1897: 716, pl. 1(13, 14); Shen 1940a: 235; Davie 1992a: 347 [key]; Komai et al. 1995: 105; Bravo et al. 2021: tab. 2. Taxonomy: Tesch (1915), Barnes (1970), Davie (2012), Shih et al. (2015). Type: Syntypes in NHM 1886.52 (NHM Data Portal). Type locality: Mergui Archipelago. Distribution: Eastern Indian Ocean to West Pacific: Bay of Bengal, Peninsular Malaysia, Gulf of page 113 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Thailand, South China and Taiwan, to Indonesia and northern Australia (Davie 2012; Shih et al. 2015). Habitat: Intertidal; substrates of fine sand or mud, near or under shades of mangroves. Macrophthalmus latreillei (Desmarest, 1822) # Macrophthalmus serratus — Stimpson 1858b: 97; Stimpson 1907: 96, pl. 13(3); Gee 1926: 165 [list]. Macrophthalmus latreillei — Gordon 1931: 528; Shen 1940a: 234; Morton and Morton 1983: 157, fig. 11.12(4); Ong Che and Morton 1991: tab. 1; Davie 1992a: 348 [key]; Davie 1992b: tab. 1; SY Lee and Leung 1999: 73; Wong et al. 2021: 66, fig. 103, pls. 19F, 20A. Macrophthalmus (Venitus) latreillei — Barnes 1970: 236. Macrophthalmus latreille — Bones 1982: 691, pl. 3c, d; Morton and Scott 1989: 140, fig. 11. Macrophthalmus latreilli — Blackmore and Rainbow 2000: app. 1. Venitus latreillei — Bravo et al. 2021: tab. 2. Taxonomy: Barnes (1967 2010), Davie (2012), Shih et al. (2015). Type: Syntypes probably in MNHN (Davie 2002). Type locality: “Indes orientales”. Distribution: Western Indian Ocean to West Pacific: from South Africa, Madagascar, to India, Southeast and East Asia to Japan, New Caledonia and Australia (Davie 2012; Shih et al. 2015). Habitat: Intertidal and shallow subtidal; substrates of soft mud in estuaries. Remarks: Although M. serratus Adams & White, 1849 was still listed as a valid taxon (as in Ng et al. 2008), this species has been regarded a junior synonym of M. latreillei (Desmarest, 1822) (Barnes 1967 1977) or “... as being synonyms of species within this group” (p. 44, Barnes 2010). The local record by Stimpson (1858b 1907) is thus placed under that of the better understood M. latreillei. We follow Davie (2012) in recognizing this species under the genus Macrophthalmus. Macrophthalmus pacificus Dana, 1851 # Macrophthalmus pacificus — Gordon 1931: 528; Shen 1940a: 235; Davie 1992a: 348 [key]; Bravo et al. 2021: tab. 2. Macrophthalmus (Mareotis) pacificus — Barnes 1970: 232. Taxonomy: Barnes (1967), Komai et al. (1995), Davie (2012). Type: Presumably lost. Type locality: Upolu, Samoa. Distribution: Arabian Sea to West Pacific: Western India to Phuket (Thailand), Southeast and East Asia reaching Ryukyus (Japan), and New Guinea, northwestern to eastern Australia, Solomon Islands and Samoa (Komai et al. 1995; Davie 2012). Habitat: Supratidal to intertidal: on mudflats under shades of mangroves, with substantial freshwater influences. Macrophthalmus tomentosus Eydoux & Souleyet, 1842 # Macrophthalmus (Mareotis) tomentosus — Davie 1992a: 346, pl. 1A, B; SY Lee and Leung 1999: 73; Bravo et al. 2021: tab. 2. Taxonomy: Barnes (1970), Komai et al. (1995), Shih et al. (2015). Type: Syntype probably in MNHN. Type locality: Manila, the Philippines. Distribution: Eastern Indian Ocean to West Pacific: from Mergui Archipelago, Phuket (Thailand), Peninsular Malaysia, the Philippines, South China, Taiwan, to Indonesia and New Caledonia (Shih et al. 2015). Habitat: Intertidal; on mudflats of very soft substrates with some freshwater influences. Subfamily Tritodynamiinae Števčić, 2005 Tritodynamia dilatata Yang & Sun, 1996 # Tritodynamia dilatata — Wong et al. 2021: 66, fig. 104, pl. 20B; Ng and Ho 2023: 210. Taxonomy: Yang and Sun (1996), Naruse and Ng (2010). Type: Holotype ♂ BNHM-J96037. Type locality: Off east coast of Guangdong, China. Distribution: South China: Guangdong and Fujian. Habitat: Subtidal; on sandy substrates of depths less than 20 m. Remarks: Ng and Ho (2023: 210) discussed the taxonomy of T. dilatata and T. longipropoda Dai, in Dai, Feng, Song & Cheng, 1980, both of which were described from Guangdong in southern China and suggested that they may be synonymous. The two species can only be separated by differences in the ambulatory leg proportions, and they noted that other characters used by Dai et al. (1980) T. longipropoda from T. dilatata are subjected to variation. The material from Hong Kong examined by Ng and Ho (2023: 201) was incorrectly listed under T. longipropoda by these authors. Tritodynamia horvathi Nobili, 1905 Tritodynamia horvathi — Morton and Morton 1983: 204, fig. 9.13(6a). Taxonomy: Shen (1932b), Sakai (1939). Type: Syntypes in HNHM and MRSN. page 114 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Type locality: Kobe, Japan. Distribution: East Asia: North China, Korea and Japan (Yang et al. 2008). Habitat: Intertidal to subtidal; commensal with polychaete worm Loimia medusa (Savigny, in Lamarck, 1818) (Morton and Morton 1983). Tritodynamia rathbunae Shen, 1932 Tritodynamia rathbuni — Morton and Morton 1983: 204, fig. 9.13(3a). Taxonomy: Shen (1932b), Sakai (1939). Type: Holotype ♂ ZMFMIB No. 4913: status unknown. Type locality: Yantai (as Chefoo), Shandong, China. Distribution: East Asia: coasts of China, Korea, and Japan (Yang et al. 2008). Habitat: Intertidal to subtidal; commensal with polychaete worm Mesochaetopterus japonicus Fujiwara, 1934 (Morton and Morton 1983). Family Mictyridae Dana, 1851 Mictyris brevidactylus Stimpson, 1858 # Mictyris longicarpus — Ortmann 1894: 748 [part: Hong Kong material only]; Gordon 1931: 528; Shen 1940a: 231; Morton and Morton 1983: 153, fig. 9.4(6); Tam and Wong 2000: 116, 2 unnumb. figs.; BKK Chan and Caley 2003: 88, 2 unnumb. figs.; Fong et al. 2005: 51, 11 unnumb. figs. Mictyris brevidactylus — Stimpson 1907: 103, pl. 13(5) [part: Hong Kong material only]; Davie et al. 2010: 94, figs. 2B, 3, 4G, H. Myctiris brevidactylus — Gee 1926: 165 [list]; Bravo et al. 2021: tab. 2. Dotilla myctiroides — Hill et al. 1975: 65; Hill et al. 1978: 92; Morton 1979a: 118, fig. 6.18r. Myctris longicarpus — Morton 1976: 102, fig. 55c. Dotilla mycteroides — Hill and Phillipps 1981: 152, pl. 51c. Taxonomy: Takeda (1978), Davie et al. (2010). Type: Neotype ♂ WAM-237-80 (designated by Davie et al. 2010). Type locality: Tong Fuk, Lantau, Hong Kong (neotype). Distribution: South China Sea and East Asia: northern Vietnam, South China and Taiwan. Habitat: Intertidal; on sheltered sandy shores or mudflats. Remarks: The soldier crab M. brevidactylus Stimpson, 1858 in East Asia has long been in synonymy M. longicarpus Latreille, 1806, but the latter species is now recognized as being present only in the Southern Hemisphere (Takeda 1978). Stimpson’s (1858b 1907) Mictyris brevidactylus is based on material from Hong Kong (South China) and Ryukyus, Japan. Specimens from the Ryukyus have been recognized as distinct and described as M. guinotae Davie, Shih & Chan, 2010 (Davie et al. 2010). Family Ocypodidae Rafinesque, 1815 Subfamily Gelasiminae Miers, 1886 Remarks: Traditional understanding of distinct lineages of ghost crabs (“Ocypodinae”) and fiddler crabs (“Ucinae”) has been challenged, and “fiddler crabs” (or Uca sensu lato) have been shown to be paraphyletic (Shih et al. 2016a). Revised classification presented in Shih et al. (2016a), showing Ocypodinae Rafinesque, 1815 now containing ghost crab Ocypode Fabricius, 1798 (circumtropical), Uca Leach, 1814 sensu stricto (Americas) and Afruca Crane, 1975 (eastern Atlantic region) (the latter two being fiddler crab genera), and Gelasiminae Miers, 1886 accommodating all other fiddler crab genera, under which all subgeneric ranks raised to full generic status, is herein followed. Austruca lactea (De Haan, 1835) # Gelasimus lacteus — Stimpson 1858b: 100; Stimpson 1907: 108; Shen 1940a: 231. Gelasimus lactens — Koelbel 1897: 717. Uca lactea — Gordon 1931: 528; Hill et al. 1975: 65; Morton 1976: 102, fig. 55g; Hill et al. 1978: 92; Morton 1979a: 118, figs. 6.18o, 6.19; Morton and Morton 1983: 153, fig. 10.3(4); Choi 1992: 901; Jones and Morton 1994: 28, fig. 6A–I, pls. 2G, H, 3E, F; BKK Chan and Caley 2003: 92, 2 unnumb. figs.; Fong et al. 2005: 56, 1 unnumb. fig.; WPW Kwok and Tang 2006: 4, figs. 10, 11; So and Lui 2007: 31, 5 unnumb. figs.; Shih et al. 2010b: 10. Uca (Celuca) lactea lactea — Crane 1975: 300, figs. 19A, 54J, JJ, 69E, pl. 40A, B. Uca (Celuca) lactea — SY Lee and Leung 1999: 71. Uca lactea annulipes — Tam and Wong 2000: 117, 2 unnumb. figs. Austruca lactea — Bravo et al. 2021: tab. 2. Taxonomy: Sakai (1939: as Gelasimus), Crane (1975: as U. (Celuca) lactea lactea), Naderloo et al. (2010: as U. (Austruca)), Shih et al. (2015). Type: Lectotype ♂ RMNH D 254, and paralectotypes in RMNH, SMF, ZMG, NHMD, MNHN (designated by Crane 1975; see also Yamaguchi and Baba 1993, Fransen et al. 1997, Naderloo et al. 2010). Type locality: Japan, locality unspecified. Distribution: South China Sea and East Asia: northern Vietnam, South China, Taiwan, Korea and Japan (Shih et al. 2010b 2022). Habitat: Intertidal; in burrows on mudflats, preferring coarser substrates. page 115 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Gelasimus borealis (Crane, 1975) # Gelasimus vocans — Stimpson 1858b: 99 [part: Hong Kong material only]; Stimpson 1907: 104 [part: Hong Kong material only]; Koelbel 1897: 717. [not Cancer vocans Linnaeus, 1758]. Uca marionis — Gee 1926: 165 [list]; Morton 1979a: 118, fig. 6.17. Uca marionis var. nitidus — Gordon 1931: 528. Gelasimus marionis — Shen 1940a: 232. Gelasimus marionis nitidus — Shen 1940a: 232. Uca (Thalassuca) vocans borealis Crane, 1975: 90, fig. 64A; SY Lee and Leung 1999: 71. Uca vocans — Hill et al. 1975: 48; Morton 1976: 102; Hill et al. 1978: 92; Morton 1979a: 121; Melville and Morton 1983: 11, 1 unnumb. fig. (13); Morton and Morton 1983: 184, fig. 11.11(5). [not Cancer vocans Linnaeus, 1758]. Uca gaimardi — Hill et al. 1975: 77; Hill et al. 1978: 107; Hill and Phillipps 1981: 152, pl. 51a. [not Gelasimus Gaimardi H. Milne Edwards, 1852 = G. crassipes White, 1847]. Uca gaimardii — Morton 1976: 103. [not Gelasimus Gaimardi H. Milne Edwards, 1852 = G. crassipes White, 1847]. Uca borealis — Jones and Morton 1994: 22, fig. 4A–I, pls. 2E, F, 3A, B; BKK Chan and Caley 2003: 92, 3 unnumb. figs.; Fong et al. 2005: 56, 1 unnumb. fig.; WPW Kwok and Tang 2006: 4, figs. 6, 7; So and Lui 2007: 30, 4 unnumb. figs.; Shih et al. 2010a: 52; 2010b: 8. Uca vocans vocans — Tam and Wong 2000: 118, 2 unnumb. figs. [not Cancer vocans Linnaeus, 1758]. Gelasimus borealis — Bravo et al. 2021: tab. 2. Taxonomy: Crane (1975: as U. (Thalassuca) vocans borealis), Shih et al. (2015). Type: Holotype ♂ USNM 137669, and paratypes in USNM. Type locality: Hong Kong. Distribution: South China Sea and East Asia: northern Vietnam, South China, Taiwan and Japan (Shih et al. 2010b 2022). Habitat: Intertidal; on mudflats, prefers coarse or gritty sand. Remarks: So far seven species have been recognized in the “Uca vocans species complex” (Crane 1975; Shih et al. 2010a; Rosenberg 2019), while only two have been reliably reported from coasts of South China—the locally common G. borealis from Guangxi, Hainan to Fujian, whereas G. vocans sensu stricto ascertained from Hainan, with northern distributinal limit unclear (Shih et al. 2010b). As such, older records of vocans, including those by Stimpson and Koelbel, might instead represent, or at least include the locally common G. borealis. Gelasimus marionis Desmarest, 1823 was indicated by Holthuis (1959) to be a junior synonym of Cancer vocans Linnaeus, 1758. Shen (1940a) reported two distinct forms, namely G. marionis and G. m. nitidus, both citing the work by Alcock (1900b). In the dichotomous key provided by Alcock, these two forms are distinguished by form of crest on internal surface of male cheliped, and shape of pollex along cutting margin (p. 353, Alcock 1900b). This distinction, however, reflects simultaneous brachychelous and leptochelous conditions commonly found in a single population of Gelasimus vocans sensu lato, and we place both entries under G. borealis. As for “U. gaimardii”, this name is now considered synonymous with Paraleptuca crassipes (Crane 1975). However, Hill and Phillipps (1981, pl. 51a) presented a peculiar illustration under this name that seems to show features of two fiddler crab species: the form of male cheliped, externally tubercular and pollex scalloped, corresponds with G. borealis, but the grayish carapace bearing dark transverse bands (discernable in the printed copy of their book), resembles P. splendida. We place this record under G. borealis herein. Paraleptuca splendida (Stimpson, 1858) # Gelasimus splendidus Stimpson, 1858b: 99; Alcock 1900b: 355; Stimpson 1907: pl. 14(2). Uca splendida — Gee 1926: 165 [list]. Uca gaimardi — Gordon 1931: 528. Gelasimus gaimardi — Shen 1940a: 232. Uca (Amphiuca) chlorophthalmus crassipes — Crane 1975: 101 [part: Hong Kong material only], figs. 13G, 14B, BB; SY Lee and Leung 1999: 71. Uca chlorophthalmus crassipes — Choi 1992: 901; Tam and Wong 2000: 117, 2 unnumb. figs. Uca chlorophthalmus — Morton and Morton 1983: 184, fig. 10.3(3). Uca chlorophthalmus — SY Lee 1993: tab. 1. Uca crassipes — Jones and Morton 1994: 26, fig. 5A–J, pl. 3C, D; BKK Chan and Caley 2003: 92, 2 unnumb. figs.; Fong et al. 2005: 57, 1 unnumb. figs.; WPW Kwok and Tang 2006: 4, figs. 8, 9; So and Lui 2007: 32, 4 unnumb. figs.; Shih et al. 2010b: 9. Uca chloropthamalus — BKK Chan 2001: 24, fig. 1b. Uca (Paraleptuca) splendida — Shih et al. 2012: 34, figs. 2–4, 6, 7C. Paraleptuca splendida — Bravo et al. 2021: tab. 2. Taxonomy: Shih et al. (2012: as Uca (Paraleptuca); 2015). Type: Neotype ♂ ZRC 2012.0143 (designated by Shih et al. 2012). Type locality: Tai Tam, Hong Kong (neotype). Distribution: South China Sea and East Asia: Central Vietnam to South China, Taiwan, and Ryukyus (Japan) (Shih et al. 2012 2016b 2022). Habitat: Supratidal to intertidal; in burrows on salt marshes, substrates of coarse sands and rock fragments, rather compact, covered by coastal vegetation. Remarks: Crane (1975) recognized two distinct subspecies in her Uca (Amphiuca) chlorophthalmus: chlorophthalmus from the western Indian Ocean along eastern Africa, and crassipes from the western and central Pacific (Shih et al. 2013). Gelasimus gaimardi H. Milne Edwards, 1852 is a junior synonym of G. crassipes White, 1847 according to Crane (1975). Among material identified as crassipes, Crane (1975) page 116 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan noted in some larger specimens from Hong Kong, the morphology of distal tip of the G1 was aberrant being “strikingly longer”, and suggested G. splendidus Stimpson, 1858 “would certainly have been kept to designate at least a subspecies” (p. 99). She, however, observed too much variation, especially in smaller specimens, to be sure. Stimpson’s (1858b) species was eventually resurrected by Shih et al. (2012) based on morphological and molecular approaches. Tubuca acuta (Stimpson, 1858) # Uca mani — Gordon 1931: 528. Gelasimus manii — Shen 1940a: 232. Gelasimus urvillei — Shen 1940a: 232. Uca (Deltuca) acuta acuta — Crane 1975: 25, fig. 61B, pl. 1E–H. Uca acuta — SY Lee 1993: tab. 1; Jones and Morton 1994: 12, fig. 1A–K, pls. 1A, B, 2A, B; So and Lui 2007: 27, 3 unnumb. figs.; Shih et al. 2010b: 3, fig. 2A–D. Uca (Deltuca) acuta — SY Lee and Leung 1999: 70. Tubuca acuta — Bravo et al. 2021: tab. 2. Taxonomy: Crane (1975: as U. (Deltuca) acuta acuta), Shih et al. (2010b: as Uca; 2015). Type: Neotype ♂ USNM 137665 (designated by Crane 1975). Type locality: Castle Peak area, Kowloon, Hong Kong (neotype). Distribution: South China Sea and East Asia: northern Vietnam, South China, and Taiwan (Shih et al. 2015 2022). Habitat: Intertidal; on estuarine mudflats, often associated with vegetation. Remarks: Uca manii Rathbun, 1909 was a name given to replace the record of Gelasimus acutus reported from the Mergui Islands by De Man (1888b), which later synonymized under Gelasimus forcipatus Adams and White, 1849 by Crane (1975). Gelasimus urvillei H. Milne Edwards, 1852 is shown to be an eastern African species (Shih et al. 2018), despite type locality listed “Vanikoro” (= now Vanuatu), which probably erroneous (Crane 1975). Both species are unlikely to be present in Hong Kong, and we follow Shih et al. (2010b) in considering these as U. acuta (now Tubuca). Tubuca arcuata (De Haan, 1835) # Gelasimus arcuatus — Shen 1940a: 231. Uca (Deltuca) arcuata — Crane 1975: 44, figs. 8C, 9C, 61J, pl. 5A–F; SY Lee and Leung 1999: 70. Uca arcuata — Melville and Morton 1983: 11, 1 unnumb. fig. (13); Morton and Morton 1983: 184, fig. 11.11(6), pl. 19H; SY Lee 1993: tab. 1; Jones and Morton 1994: 13, fig. 2A–J, pls. 1C, D, 2C, D; Tam and Wong 2000: 116, 2 unnumb. figs.; Fong et al. 2005: 56, 1 unnumb. fig.; WPW Kwok and Tang 2006: 4, figs. 4, 5; So and Lui 2007: 28, 3 unnumb. figs.; Shih et al. 2010b: 7, fig. 2E, F. Uca acuta — WPW Kwok and Tang 2006: 4, fig. 2. Tubuca arcuata — Bravo et al. 2021: tab. 2. Taxonomy: Shen (1932b), Crane (1975: as U. (Deltuca) arcuata), Shih et al. (2015 2016b). Type: Lectotype ♂ RMNH D 243, and paralectotypes in RMNH, NHM, NHMD, MNHN (designated by Crane 1975; see also Yamaguchi and Baba 1993; Fransen et al. 1997). Type locality: Japan, locality unspecified. Distribution: South China Sea and East Asia: northern Vietnam, coasts of China, Taiwan, Korea and Japan (Shih et al. 2015 2016b 2022). Habitat: Intertidal; on estuarine mudflats, substrates of soft mud. Tubuca coarctata (H. Milne Edwards, 1852) Tubuca coarctata — Jimenez et al. 2024: 182, figs. 3A, B, 4A, B, 5. Taxonomy: Crane (1975: as U. (Deltuca) coarctata coarctata), Shih et al. (2015 2016b). Type: Lectotype ♂ and paralectotypes in MNHN (designated by Crane 1975). Type locality: Odessa, Ukraine (possible mislabeling; Crane 1975). Distribution: West Pacific: the Philippines, Indonesia, to South China, Taiwan and Okinawa, to New Guinea and northern and northeastern Australia (Crane 1975; Shih et al. 2015 2016b). Tubuca dussumieri (H. Milne Edwards, 1852) ? Gelasimus dussumieri — Shen 1940a: 232 [no new specimens]. Uca (Deltuca) dussumieri dussumieri — Dai et al. 1986: 421, fig. 233(1–4). Uca (Deltuca) dussumieri — Dai and Yang 1991: 461, fig. 233(1–4). Uca dussumieri — Shih et al. 2010b: 9 [no new specimens]. Tubuca dussumieri — Jimenez et al. 2024: 182, figs. 3C–H, 4C–F, 6. Taxonomy: Crane (1975: as U. (Deltuca) dussumieri dussumieri), Shih et al. (2015). Type: Lectotype ♂ in MNHN (designated by Crane 1975). Type locality: Samarang, Java, Indonesia. Distribution: West Pacific: New Guinea, Indonesia, Philippines, South China (Hong Kong), Taiwan and Ryukyus (Crane 1975; Shih et al. 2016b). Habitat: Intertidal; on muddy substrates with coarse sediments, sometimes near shades of vegetation (Shih et al. 2015). Remarks: Despite report from “Guangdong” by Dai et al. (1986) and Dai and Yang (1991), presences of this species along shores of South China remained page 117 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan unverified (Shih et al. 2010b). However, as pointed out by the late Prof. S.-L. Yang (pers. comm. to HTS), the illustrated specimen from “Guangdong” was actually collected from Hong Kong, with no precise date or data otherwise known (see Shih et al. 2010b). Throughout excursions spanning the 2010s, no occurrence of this species was encountered. Recent investigations, however, show the species to occur locally in small populations (Jimenez et al. 2024), and herein included. Tubuca paradussumieri (Bott, 1973) # ? Uca dusummieri — Hill et al. 1975: 77; Morton 1976: 105; Hill et al. 1978: 107; Morton 1979a: 121; SY Lee 1993: tab. 1. Uca paradussumieri — Jones and Morton 1994: 18, fig. 3A–H, pl. 1E, F; WPW Kwok and Tang 2006: 4, figs. 1, 12, 13; So and Lui 2007: 29, 4 unnumb. figs.; Shih et al. 2010b: 10, fig. 3A–D. Uca (Deltuca) paradussumieri — SY Lee and Leung 1999: 70. Uca acuta — WPW Kwok and Tang 2006: 4, fig. 3. Tubuca paradussumieri — Bravo et al. 2021: tab. 2. Taxonomy: Crane (1975: as U. (Deltuca) dussumieri spinata), Shih et al. (2010b: as Uca; 2015). Type: Holotype ♂ in SMF 5650, paratypes in SMF, NHMD, ZSM. Type locality: Bengal. Distribution: Eastern Indian Ocean to South China Sea: from Bay of Bengal, Thailand, Peninsular Malaysia, Borneo, Sumatra (Indonesia), to Vietnam, South China and Taiwan (Shih et al. 2015 2022). Habitat: Intertidal; on estuarine mudflats, substrates of soft mud, often under shades of mangroves. Subfamily Ocypodinae Rafinesque, 1815 Ocypode ceratophthalmus (Pallas, 1772) # Ocypode ceratophthalma — Stimpson 1858b: 100; Stimpson 1907: 108, pl. 7(2); George 1982: 187, pl. 2; Morton and Morton 1983: 141, fig. 9.4(1, 2); ECK Wong 1990: 150; Choi 1992: 901; BKK Chan and Caley 2003: 90, figs. 1–3; BKK Chan et al. 2006: 44, fig. 1a. Ocypoda ceratophthalma — Gordon 1931: 528; Shen 1940a: 232. Ocypode gaimardi Hill, Gott & Morton, 1975: 65 [nomen nudum]; Hill, Gott, Morton & Hodgkiss, 1978: 92 [nomen nudum]; Morton 1976: 101; Morton 1979a: 118; Hill and Phillipps 1981: 152, pl. 51d. Ocypode ceratophthalmus — Hill et al. 1975: 65; Morton 1976: 101; Hill et al. 1978: 92; Morton 1979a: 118, fig. 6.18e; Hill and Phillipps 1981: 152, pl. 51e; Fong et al. 2005: 53, 5 unnumb. figs.; K Sakai and Türkay 2013: 685, figs. 1D–I, 10, 32; Bravo et al. 2021: tab. 2. Taxonomy: George and Knott (1965), K Sakai and Türkay (2013), Shih et al. (2015). Type: Whereabouts and status unknown. Type locality: India (George and Knott 1965). Distribution: Western Indian Ocean to East Pacific: from South Africa, Madagascar, India, Southeast and East Asia, to Australia and New Caledonia, towards Samoa, Hawaii and Clipperton Island (K Sakai and Türkay 2013). Habitat: Intertidal; in deep tube-like burrows along exposed sandy shores (BKK Chan et al. 2006). Remarks: We are unable to trace identities of the name “Ocypode gaimardi” as presented in Hill et al. (1975 1978), Morton (1976 1979a) and Hill and Phillipps (1981) elsewhere in the literature. In the latter work, a guidebook, color illustrations under this name (pl. 51d) clearly depict a juvenile ghost crab undoubtedly identifiable as O. ceratophthalmus. Throughout the vast geographical distribution of this species, based on sequences of the mitochondrial COI marker, Ma et al. (2018) recently showed this species shows substantial genetic structuring in the Indo-Pacific. There are three major genetically distinct clades: east, central and west clades in the Indo-Pacific (Ma et al. 2018). The population in Hong Kong belongs to the central clade as presented by Ma et al. (2018). B.K.K. Chan and colleagues (2006) investigated burrow morphologies of juvenile and adults of O. certatophthalmus in Hong Kong. Ocypode mortoni George, 1982 # ? Ocypoda stimpsoni — Shen 1940a: 233. ? Ocypode stimpsoni — Hill et al. 1975: 65; Hill et al. 1978: 92; Morton 1979a: 118, fig. 6.17. Ocypode mortoni George, 1982: 187, figs. 1C, 2C, pl. 3; Morton and Morton 1983: 141, fig. 8.7(3); Wong et al. 2012: 75, figs. 3–6; K Sakai and Türkay 2013: 722, figs. 3E, 20, 42. Taxonomy: George (1982), Wong et al. (2012), K Sakai and Türkay (2013). Type: Holotype ♂ WAM 2-81, and paratypes in WAM. Type locality: Sai Wan, eastern new Territories, Hong Kong. Distribution: East Asia: South China and southern Japan (K Sakai and Türkay 2013). Habitat: Intertidal; in burrows along exposed sandy shores. Remarks: Ocypode stimpsoni Ortmann, 1897(b) and O. mortoni are morphologically close, identified primarily by the latter possessing ocular stylets in adults (Wong et al. 2012; K Sakai and Türkay 2013), and live coloration — O. stimpsoni dark orange-reddish when undisturbed, whereas O. mortoni is always a dull yellow and whitish in life. Along coasts of China, the southern distributional limit of the temperate O. stimpsoni had been recorded to be Zhejiang (Y Chen 1991). Although page 118 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan we cannot exclude the possibility that O. stimpsoni is or was present in Hong Kong, we are more inclined to suspect that Shen, in examining material from Hong Kong (1940a) and earlier from Hainan (1936c), had confounded the tropical form, later described as O. mortoni, under O. stimpsoni. We have surveyed local sandy shores for the past decade, but no O. stimpsoni has ever been reliably encountered. Moreover, it is difficult to believe that a conspicuous reddish Ocypode crab would have been missed by naturalists and locals. As such, we exclude it from the fauna of Hong Kong. Ocypode sinensis Dai, Song & Yang, 1985 # Ocypode cordimana — Stimpson 1858b: 100 [part]; Stimpson 1907: 110, pl. 15(2); Gee 1926: 165 [list]; George 1982: 187, pl. 1. [not fig. 2A]; Morton and Morton 1983: 141; Choi 1992: 901; BKK Chan and Caley 2003: 90, fig. 4. [not O. cordimanus Latreille, 1818]. Ocypoda cordimana — Balss 1922b: 142 [part]; Shen 1940a: 232. Ocypode sinensis Dai, Song & Yang, 1985: 372, figs. 8–12, pl. 1(3–6); J-F Huang et al. 1998: 943, figs. 1a, 2a, c, e, g, 3a, c, e, f, 4a, c–e; Bravo et al. 2021: tab. 2. Taxonomy: Dai et al. (1985), J-F Huang et al. (1998), Shih et al. (2015). Type: Holotype ♂ BNHM J97088b, and paratypes in BNHM and IZAS (= IZCAS) (J-F Huang et al. 1998). Type locality: Jinyindao, Xisha (= Paracel) Islands, South China Sea. Distribution: Eastern Indian Ocean to South China Sea and East Asia: East India, Christmas Island, Peninsular Malaysia, Borneo, the Philippines, South China, Taiwan and Japan. Habitat: Supratidal; in deep burrows along exposed sandy shores, often among low coastal vegetation. Remarks: The widespread O. cordimanus Latreille, 1818 is repeatedly reported throughout the entire IndoWest Pacific region. Older local records under this name had probably included (or entirely representing) another similar species later described from South China as O. sinensis Dai, Song & Yang, 1985. K. Sakai and Türkay (2013) treated O. sinensis merely as a juvenile of O. cordimanus but the available morphological and molecular evidence indicates both are distinct species (J-F Huang et al. 1998; Wong et al. 2012; Shih et al. 2015). We have no reliable reports of O. cordimanus sensu stricto in Hong Kong as yet. Family Xenophthalmidae Stimpson, 1858 Subfamily Anomalifrontinae Rathbun, 1931 Anomalifrons lightana Rathbun, 1931 # (Figs. 34, 41D) Anomalifrons lightana — Khot et al. 2019: 1343. Taxonomy: Rathbun (1931), Serène and Umali (1972), Khot et al. (2019). Type: Holotype ♂ USNM 61878. Type locality: Guantou, near Fuzhou, Fujian, China. Distribution: Southeast and East Asia: Peninsular Malaysia, northern Vietnam to South China (Rathbun 1931; Serène and Umali 1972; Wada 2019). Habitat: Intertidal to subtidal; on sandy-muddy substrates, with substantial freshwater infleunces. Subfamily Xenophthalminae Stimpson, 1858 Neoxenophthalmus obscurus (Henderson, 1893) # Neoxenophthalmus obscurus — Shin 2000: tab. 1; Shin et al. 2003: tab. 3; Shin et al. 2004: 130, tab. 2; Wong et al. 2021: 66, fig. 105, pl. 20C. Taxonomy: Rathbun (1910: as Xenophthalmus), Serène and Umali (1972). Type: Holotype ♀ BM 88.34 in NHM (Sankarankutty 1969; Schmitt et al. 1973). Type locality: Gulf of Martaban, Myanmar. Distribution: Eastern Indian Ocean to South China Sea: East India, Myanmar, Thailand, South China and Taiwan. Habitat: Subtidal; on sandy-muddy substrates at shallow depths. Xenophthalmus pinnotheroides White, 1846 # Xenophthalmus pinnotheroides — Stimpson 1858b: 107; Stimpson 1907: 141; Gee 1926: 164 [list]; Shen 1940a: 230; Ong Che and Morton 1991: tab. 1; Shin 2000: tab. 1; Wong et al. 2021: 66, fig. 106, pl. 20D. Taxonomy: Rathbun (1910), Shen (1937a), Serène and Umali (1972). Type: Syntypes BM 43.6 (Sankarankutty 1969; Schmitt et al. 1973); NHM White 3 759. a-c (NHM Data Portal). Type locality: The Philippines. Distribution: Eastern Indian Ocean to West Pacific: southeastern India, Thailand, the Philippines, China, Japan, Indonesia and Queensland (Australia) (Schmitt et al. 1973). page 119 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Habitat: Subtidal; substrates sandy-muddy at shallow depths. Superfamily Pinnotheroidea De Haan, 1833 Family Pinnotheridae De Haan, 1833 Subfamily Pinnothereliinae Alcock, 1900 Indopinnixa mortoni Davie, 1992 Indopinnixa mortoni Davie, 1992a: 354, fig. 1A–G. Taxonomy: Davie (1992a). Type: Holotype ♂ QM W16480, and paratype ♀ in QM. Type locality: Hoi Ha Wan, New Territories, Hong Kong. Distribution: So far South China only. Habitat: Intertidal; in burrows of sandy-mud substrates between rocks, commensal in mud tubes of large capitellid polychaetes (Davie 1992a). Indopinnixa sipunculana Manning & Morton, 1987 Pinnixa rathbuni — Morton and Morton 1983: 78. Indopinnixa sipunculana Manning & Morton, 1987: 544, figs. 1, 3C; Morton 1988: 40, pl. 9. Taxonomy: Manning and Morton (1987). Type: Holotype ♂ USNM 221697, and paratype ♀ in USNM. Type locality: Tai Tam, Hong Kong. Distribution: So far from Hong Kong only. Habitat: Intertidal; on mudflats, commensal in burrows of the sipuculan worm Sipunculus nudus Linnaeus, 1766 (Manning and Morton 1987). Pinnixa balanoglossana Sakai, 1934 Pinnixa balanoglossana — Morton and Morton 1983: 204, fig. 9.13(5a). Taxonomy: Sakai (1934 1939). Type: Whereabouts and status unknown. Fig. 34. Anomalifrons lightana Rathbun, 1931 (male, 12.7 × 8.9 mm, Tung Chung): A, carapace; B, frontal region, ventral view; C, maxilliped 3; D, right chela; E, left P4; F, thoracic sternites and pleon; G, left G1, mesial view; H, tip of left G1, mesial view. Structures A to C denuded. page 120 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Type locality: Momotori, Ise Bay, Japan. Distribution: East Asia: endemic to Japan (Sakai 1976; Watanabe 2012); probably also in Hong Kong (Morton and Morton 1983). Habitat: Intertidal; on mudflats, commensal with acorn worm Balanoglossus misakiensis Kuwano, 1902 (Sakai 1934; Morton and Morton 1983). Pinnixa penultipedalis Stimpson, 1858 Pinnixa penultipedalis Stimpson, 1858b: 54; Stimpson 1907: 143; Gee 1926: 163 [list]; Shen 1940a: 230. Taxonomy: Stimpson (1907), Sakai (1976), Manning and Morton (1987), Naruse and Maenosono (2012). Type: Probably lost. Type locality: Hong Kong. Distribution: East Asia: probably only South China (Hong Kong) (see Remarks below). Habitat: Subtidal; among dead shells on muddy substrates in 10 fathoms (about 18.3 m) (Stimpson 1907). Remarks: The present species was described from a single female specimen reported by Stimpson (1858b), as having a very broad carapace (2.11 times as broad as long), the dorsal surface being smooth and glossy, and posterior extremity with transverse ridge. The species, however, has never been figured and the types are almost certainly lost. Ortmann (1894: pl. 23(7)) and Shen (1937a: text-fig. 10a) subsequently reported the species from Nagasaki, Japan, and Tsangkou (near Qingdao), Shandong, China, respectively. Both records are considered doubtful: the Japanese record by Ortmann (1894) has a relatively narrower carapace, whereas the record from North China, as indicated in Manning and Morton (1987), apart from a very pronounced transverse cardiac ridge, lacks any transverse ridge on posterior part of the carapace. A recent report of one male specimen from Oki Islands, Sea of Japan by Yamaguchi and Konishi (2004: fig. 1A) shows a specimen with a carapace width to length ratio of approximately 1.95 (4.18 mm × 2.14 mm) and is narrower than in the original description. As suggested by Naruse and Maenosono (2012), we believe none of these records actually represent P. penultipedalis sensu stricto. Subfamily Pinnotherinae De Haan, 1833 Amusiotheres obtusidentatus (Dai, Feng, Song & Chen, 1980) Pinnotheres obtusidentatus — Pregenzer and Morton 1990: 656, fig. 4. Amusiotheres obtusidentatus — Ng and Ho 2016: 742, figs. 1–8. Taxonomy: Dai et al. (1980: as Fabia), Ng and Ho (2016), Ng and Ngo (2022). Type: Holotype ♀ IZAS CS 01546a, no. 14270 in IZCAS (Ng and Ho 2016). Type locality: Beihai, Guangxi, China. Distribution: South China Sea: southern Thailand, Vietnam, South China, and western Taiwan (Ng and Ho 2016; Ng and Ngo 2022). Habitat: Subtidal; commensal with or parasite of bivalve Amusium pleuronectes (Linnaeus, 1758) (see Ng and Ho 2016). Arcotheres atrinae (Sakai, 1939) Pinnotheres sinensis atrinae — Markham 1990: 560; Markham 1992: tab. 1 [list]; Li 2003: 153 [no new specimens]. Taxonomy: Sakai (1939: Pinnotheres sinensis atrinae), Ng et al. (2017b). Type: Whereabouts and status unknown. Type locality: Kii Peninsula, Japan. Distribution: East Asia: endemic to Japan (Schmitt et al. 1973; Sakai 1976), probably also in South China (Hong Kong) (above cited report). Habitat: Subtidal; commensal with or parasite of bivalve Atrina japonica (Reeve, 1858) on sandy substrates (Sakai 1939). Arcotheres cyclinus (Shen, 1932) Pinnotheres cyclinus — Morton and Morton 1983: 202, fig. 9.9(7A). Pinnotheres obscurus — Pregenzer and Morton 1990: 653 [part], fig. 3A, B. Taxonomy: Shen (1932b), Sakai (1939), Ng et al. (2017b). Type: Syntypes ♀♀ in IOCAS. Type locality: Xingcun (as Hsingtsun), Shandong, China. Distribution: East Asia: coasts of China, Taiwan, Korea and Japan. Habitat: Intertidal to subtidal; commensal with or parasite of bivalves, including Cyclina sinensis (Gmelin, 1791), Meretrix meretrix (Linnaeus, 1758), Barbatia virescens (Reeve, 1844), Geloina erosa (Solander, 1786), on substrates of sand and mud, or attached to rocks (Shen 1932b; Sakai 1939; Morton and Morton 1983). Remarks: Pinnotheres cyclinus Shen, 1932(b) was placed under Arcotheres Manning, 1993 by Ng et al. (2017b), a decision we follow. We also consider material reported as P. obscurus in Pregenzer and Morton (1990) included P. cyclinus. See Remarks under page 121 of 155Zoological Studies 63:38 (2024) © 2024 Academia Sinica, Taiwan Fig. 39. A–D, F, Overall habitus. A, Demania reynaudii (H. Milne Edwards, 1834) (male, 72.6 × 52.5 mm, off Clear Water Bay); B, inset, Leptodius davaoensis Ward, 1941 (male, 19.4 × 12.3 mm, Starfish Bay); C, Atergatopsis germaini A. Milne-Edwards, 1865 (male, 64.4 × 43.8 mm, off Kau Sai Chau). D, Percnon sinense Chen, 1977 (male , 28.1 × 30.2 mm, off High Island); E, F Clistocoeloma nobile B.Y. Lee, N.K. Ng & Ng, 2023 (male, 16.5 × 12.0 mm, To Gwa Peng). F preserved coloration. page 128 of 155Zoological Studies 63:38 (2024) [Document text truncated for crawler view.]