An annotated checklist of brachyuran crabs from Taiwan (Crustacea ...

ISSN 1010-16«:i 

An annotated checklist of brachyuran crabs from 

Taiwan (Crustacea: Decapoda) 

Peter K. L. Ng, Chia-Hsiang Wang, Ping-Ho Ho 

and Hsi-Te Shih 

National Taiwan Museum Special Publication Series, Number 11 

Published by the National Taiwan Museum, Taipei, Taiwan 

Republic of China 

April 2001

National Taiwan Museum Special Publication Series, No. 11, 86pp., April 2001 I 

An annotated checklist of brachyuran crabs from Taiwan (Crustacea: Decapoda) 

Contents 

mm -i 

Abstract 2 

Introduction 2 

Material and methods 3 

General notes on classification used -4 

Checklist 5 

Family Dromiidae DeHaan, 1833 5 

Family Dynomenidae Ortmann, 1892 6 

Family Homolidae De Haan, 1839 6 

Family Latreilliidae Stimpson, 1858 6 

Family RaninidaeDe Haan, 1839 6 

Subfamily Ranininae De Haan, 1839 6 

Subfamily Lyreidinae Guinot, 1993 -7 

Subfamily Notopodinae Serene andUmali, 1972 7 

Subfamily Raninoidinae Lorenthey and Beurlen, 1929 7 

Family Dorippidae MacLeay, 1838 7 

Subfamily Dorippinae MacLeay, 1838 7 

Subfamily Ethusinae Guinot, 1977 8 

Family Cyclodorippidae Ortmann, 1892 8 

Family Leucosiidae Samouelle, 1819 8 

Subfamily Leucosiinae Samouelle, 1819 -8 

Subfamily Ebaliinae Stimpson, 1871 9 

Subfamily Philyrinae Rathbun, 1937 9 

Family Calappidae DeHaan, 1833 10 

Family Matutidae DeHaan, 1833 11 

Family Majidae Samouelle, 1819 11 

Subfamily Majinae Samouelle, 1819 12 

Subfamily Epialtinae MacLeay, 1838 12 

Subfamily Inachinae MacLeay, 1838 13 

Subfamily Mithracinae MacLeay, 1838 -13 

Subfamily PisinaeDana, 1851 13 

Family Parthenopidae MacLeay, 1838 14

H National Taiwan Museum Special Publication Series, No. 11, 86pp., April 2001 

Subfamily Parthenopinae MacLeay, 1838 14 

Subfamily Cryptopodiinae Stimpson, 1871 15 

Subfamily Daldorfiinae Ng and Rodriguez, 1986 •*•• 15 

Subfamily Lambrachaeinae Stevcic, 1994 15 

Family Aethridae Dana, 1851 -15 

Family DairidaeNg and Rodriguez, 1986 15 

Family Hymenosomatidae MacLeay, 1838 15 

Family Corystidae Samouelle, 1819 15 

Family Atelecylidae Ortmann, 1893 16 

Family Cancridae Latreille, 1803 16 

Family Portunidae Rafinesque, 1815 • 16 

Subfamily Portuninae Rafinesque, 1815 16 

Subfamily Caphyrinae Paulson, 1875 * 18 

Subfamily Carupinae Paulson, 1875 18 

Subfamily Podophthalminae Dana, 1851 18 

Subfamily Polybiinae Ortmann, 1893 18 

Subfamily Thalamitinae Paulson, 1875 18 

Family Carpiliidae Ortmann, 1893 -21 

Family Eriphiidae MacLeay, 1838 -22 

Subfamily Eriphiinae MacLeay, 1838 23 

Subfamily Dacryopilumninae Serene, 1984 23 

Subfamily Menippinae Ortmann, 1893 23 

Subfamily Oziinae Dana, 1851 24 

Family Xanthidae MacLeay, 1838 -24 

Subfamily Xanthinae MacLeay, 1838 25 

Subfamily Actaeinae Alcock, 1898 26 

Subfamily Chlorodiinae Alcock, 1898 27 

Subfamily Cymoinae Alcock, 1898 27 

Subfamily Etisinae Ortmann, 1893 27 

Subfamily Euxanthinae Alcock, 1898 28 

Subfamily Kraussiinae Ng, 1993 28 

Subfamily Liomerinae Sakai, 1976 • 28 

Subfamily Polydectinae Dana, 1851 28 

Subfamily Zalasiinae Serene, 1968 29

Ng et al. : Checklist of crabs from Taiwan IE 

Subfamily Zosiminae Alcock, 1898 29 

Family Pilumnidae Samouelle, 1819 -29 

Subfamily Pilumninae Samouelle, 1819 30 

Subfamily Eumedoninae Dana, 1852 30 

Subfamily Galeninae A. Milne Edwards, 1862 31 

Subfamily Halimedinae Alcock, 1898 31 

Subfamily Rhizopinae Stimpson, 1858 31 

Subfamily Tariaochelinae Ng and Clark, 2000 31 

Family Trapeziidae Miers, 1886 31 

Subfamily Trapeziinae Miers, 1886 31 

Subfamily Domeciinae Ortmann, 1893 -32 

Family Goneplacidae MacLeay, 1838 32 

Subfamily Goneplacinae MacLeay, 1838 33 

Subfamily Carcinoplacinae H. Milne Edwards, 1852 33 

Subfamily Chasmocarcininae Serene, 1964 34 

Subfamily Euryplacinae Stimpson, 1871 34 

Subfamily Pseudoziinae Alcock, 1898 34 

Family Geryonidae Colosi, 1923 35 

Family Ocypodidae Rafinesque, 1815 35 

Subfamily Ocypodinae Rafinesque, 1815 35 

Subfamily Macrophthalminae Dana, 1851 38 

Subfamily Scopimerinae Alcock, 1900 38 

Family Camptandriidae Stimpson, 1858 39 

Family Mictyridae Dana, 1852 39 

Family Grapsidae MacLeay, 1838 40 

Subfamily Grapsinae MacLeay, 1838 40 

Subfamily Sesarminae Dana, 1851 ; 41 

Subfamily Varuninae H. Milne Edwards, 1853 44 

Family Plagusiidae Dana, 1851 47 

Family Pinnotheridae De Haan, 1833 47 

Subfamily Pinnotherinae De Haan, 1833 47 

Subfamily Asthenognathinae Stimpson, 1858 48 

Subfamily Xenophthalminae Alcock, 1900 48 

Family Cryptochiridae Paulson, 1875 48

IV National Taiwan Museum Special Publication Series, No. 11, 86 pp., April 2001 

Family Palicidae Rathbun, 1898 48 

Subfamily Palicinae Rathbun, 1898 48 

Subfamily Crossotonotinae Moosa and Serene, 1981 48 

Family Gecarcinidae MacLeay, 1838 48 

Family Potamidae Ortmann, 1896 49 

Family Parathelphusidae Alcock, 1910 52 

Endemic taxa 52 

On the fauna from Kinmen 53 

On the fauna from Tungsha Islands 54 

General discussion 54 

Acknowledgements 55 

References 56 

Appendix 1. New records from Taiwan and Tungsha Islands 74 

Appendix 2. Taiwanese localities (Chinese) and their romanizations as used in the text 77 

Appendix 3. Journal names in Chinese or Japanese and their English translation as used in the 

text 78 

Fig. 1 79 

Fig. 2 80 

Fig. 3 • 81 

Fig. 4 82 

Fig. 5 83 

Fig. 6 84 

Fig. 7 85 

Fig. 8 •. 86

National Taiwan Museum Special Publication Series, No. 11, 86 pp., April 2001 1 

An annotated checklist of brachyuran crabs from Taiwan (Crustacea: Decapoda) 

An annotated checklist of brachyuran crabs from Taiwan 

(Crustacea: Decapoda) 

Peter K. L. Ng \ Chia-Hsiang Wang 2 , Ping-Ho Ho 3 and Hsi-Te Shih *• s 

Accepted November 28,2000 

mm& £»ff¥ H4^ mm® xmm&mi&mznMzm^mm: 

+JEB) W±LWfiffi$lf£MMMsM ' Ini,86PP.,April2001 

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SIH&^JB Thacanophrys Griffin and Transfer, 1986 ' J§ PrismatopusWaid, 1933 E^ 

H4!r I H^HWilll GlobopilumnusBalss, 1933 ' Ji EupilumnusKossmann, 1877 

ft#£j^;g ; $>lf^TO55ii MopsocarcinusBarnes, 1967 (Monophthalmus M ) ' Ji 

Chaenostoma Stimpson, 1858 6*J #C H £i » ^^0H^S&*J Galene granulosa Miers, 

1884 • !§ G. bispinosa(Herbst, 1783 )#)#*£ ° 

sa&^: !£#££*' spura»mm • *fia»>#« ° 

Abstract 

An annotated checklist of the brachyuran crabs known thus far from Taiwan, together 

with citations, is compiled. The brachyuran fauna now stands at 548 species, representing 

36 families. Twenty new records are added through the present study. The most diverse 

1 

Department of Biological Sciences, National University of Singapore, Kent Ridge, Singapore 119260, 

Republic of Singapore 

2 

Department of Zoology, National Taiwan Museum, 48 Hsuchow Road, Taipei, Taiwan 100, Republic of 

China 

* National Museum of Marine Biology and Aquarium, 2 Houwan Road, Checheng, Pingtung, Taiwan 944, 

Republic of China 

* Department of Biological Sciences, National Sun Yat-sen University, 70 Lienhai Road. Kaohsiung, 

Taiwan 804, Republic of China 

3 

Current address: Department of Zoology, National Museum of Natural Science. 1 Kuanchien Road, 

Taichung, Taiwan 404, Republic of China

2 

National Taiwan Museum Special Publication Series, No. 11, 86 pp., April 2001 

family is the Xanthidae while the best studied is the Ocypodidae. A substantial proportion of 

the present records were only obtained over the last decade, with a good part of these being 

deep-water taxa. Inaddition, 47 species are now known from Kinmen (Quemoy), ofwhich 

seven are not known from Taiwan proper; and 10 species have been recorded from the Tungsha 

Islands C Pratas Islands), of which five are not known from the main island thus far. The 

species endemic to Taiwan and those now known from Kinmen and Tungsha Islands are also 

briefly discussed. It is expected that with increased sampling, the total number of brachyuran 

species will still increase substantially over the years. Several nomenclatural actions are 

also made, notably the majid genus Thacanophrys Griffin and Tranter, 1986, is a junior 

synonym of Prismatopus Ward, 1933; the eriphiid genus GlobopilumnusBalss, 1933, isa 

junior synonym of Eupilumnus Kossmann, 1877; the ocypodid subgenus Mopsocarcinus 

Barnes, 1967 (genus Macrophthalmus), is ajunior synonym of Chaenostoma Stimpson, 

1858; and the pilumnid species Galene granulosaMiers, 1884, is ajunior synonym of G. 

bispinosa (Herbst, 1783), 

Keywords: Annotated checklist, brachyuran crabs, Taiwan, newrecords, taxonomy. 

Introduction 

The brachyuran crabs of Taiwan have been well studied over the years. In particular, the reports by 

Maki and Tsuchiya (1923), Sakai (1937, 1938a, 1939), Horikawa (1940), Lin (1949), Chang 

(1963 ) and Sakai (1976 ) of the Taiwanese fauna are important. Unfortunately, only a part of their 

collection is in the National Taiwan Museum, with the whereabouts of much of the rest not known. In 

addition to these key reports, there have also been many other smaller but nevertheless important papers 

over the years on various aspects of the Taiwanese crab fauna. 

The National Taiwan Museum in Taipei is currently the oldest natural history museum in Taiwan, 

and has a substantial collection of dried and wet-preserved crab specimens, some of which date back to 

pre-Second World War days. The museum had also obtained (by purchase) two large lots of dried 

specimens from a private collector (Mr. Chun-Yang Wei) in the 1970s which had been collected between 

1950 and 1970s (see Wang and Chen, 1981). The second and third authors, as well as many of 

their colleagues, have also been collecting extensively in Taiwan over the last 10 years, and these specimens 

have been the basis of many reports (e. g. Fukui, Wada and Wang, 1989). Although much of 

the recent collections as well as those by Mr. Wei had been obtained from fishermen and trawlers, most 

are from inshore fish ports and are probably from near Taiwanese waters. It cannot be discounted, however, 

that some of the older Taiwanese records may in fact have been based on specimens originally 

collected from China, Japan, Philippines or even Southeast Asia porteeted by these boats. The present 

compilation also does not consider the fossil record, even for relatively recent ones (e. g. Hu and Tao, 

1996; but see critique in Ng, 1999a). 

The present paper lists the known extant species from Taiwan, excluding Kinmen (Quemoy) and 

Tungsha Islands (Pratas Islands). Because Kinmen and Tungsha Islands are some distance from the 

main island of Taiwan, their fauna is possibly rather different. For completeness, however, the species 

known from these areas are treated briefly in the discussion. As regards the brachyuran fauna of the po-

Ng et al. : Checklist of crabs from Taiwan 3 

litically sensitive Tiaoyutai Islands ( = Diaoyutai Islands, Senkakushato), we have decided not to treat 

its fauna. The invertebrate fauna of these islands has been documented by Nakasone and Nagahama 

(1971) with additional land crab (Gecarcinidae) records by Ng, Nakasone and Kosuge (2000). 

The present annotated list is intended to help the growing number of scientists in and outside Taiwan 

who are interested in crab biology, keeping them up to date as to what species are known, their 

current scientific names and the major publications reporting on them. As such, we have tried to keep the 

data within as current as possible. 

Material and methods 

Citations for each of the Taiwanese records are provided as far as possible. The literature cited is 

only for the Taiwanese records and author citations for individual taxa have been omitted due to space 

constraints. We had an enormous problem selecting the publications to be listed here. We had originally 

hoped to list all the papers which mention Taiwanese crabs, but this eventually proved impractical. Because 

of Taiwan's long history, there have been many small papers reporting on its fauna, particularly 

by Japanese workers (Taiwan was under Japanese rule from 1895-1945). While we have tracked down 

many of them, we have not been able to find all. Most of these papers, however, are small reports 

which are not significant. There is also an enormous amount of "grey" literature circulating in Taiwan - 

in-house journal reports, department reports, magazine articles, booklets, pamphlets, natural history 

guides etc., and these often deal with its crab fauna directly or in passing. We found many of these but 

we are sure a good number have been missed. Considering the popularity of crabs for seafood, fishing, 

aquarium, divers and photographic buffs, it would have been impossible to search for all the "grey" literature 

written by these enthusiasts. We are confident we have found (and listed) those few which contain 

useful scientific information. Papers or articles in which only the Chinese or Japanese names are cited 

(i. e. no scientific names provided) have been excluded. 

For the included literature, we have tried to ensure the taxonomic ones have all been listed. Papers 

on other biological aspects (e. g. ecology, ethology or toxicology) are listed only when the species they 

treat include new records. The same methodology is followed when popular articles, book chapters and 

general guidebooks are concerned. All books on crustaceans, however, have been included as we feel 

these contain a useful amount of scientific information. Some of the older records cannot be substantiated 

and we have used our best judgement in determining if they are accurate. In a few instances, the 

identifications are clearly incorrect but we do not know what the original authors were referring to as 

their specimens are no longer extant, and as such, we have no choice but to omit them. Although we 

have tried to identify the old records as far as possible, in cases where only generic names were provided, 

we have chosen to omit these records. This is especially the case for Lin's (1949) paper, although 

we have made one exception for his record of " Asthenognathus sp. " (Pinnotheridae), it being the only 

report of this genus from Taiwan. 

The authors also follow the general guidelines established by the International Code of Zoological 

Nomenclature (1999 ) as to what constitutes a publication (e. g. printed using conventional presses and 

distributed to a wide audience) and in many cases, these also have ISBN or ISSN numbers. On these 

criteria, some special reports (e. g. Jeng, Jan, Tzeng, Feng and Yang, 1994; Jeng, Jan, Tzeng, Feng and 

Yang, 1996; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998) have been 

included. In any case, these have been prepared or aided by competent carcinologists so the names in-

4 National Taiwan Museum Special Publication Series, No. 11, 86 pp., April 2001 

eluded are very likely to be accurate. However, we have decided to omit some of these reports because 

they have been superceded by equivalent updated ones (e. g. Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 

1997). Mention should, however, be made of the compilation by Hu and Tao (1996) of the fossil crustacean 

fauna of Taiwan, in which these authors endeavour to show what extant species are also present. 

However, they do not always indicate where their extant specimens were collected from; and many of 

their identifications are very suspicious (see Ng, 1999a). As such, we have decided, albeit reluctantly, 

to omit the records from this book for fear of causing unnecessary confusion. In any case, there are no 

obvious new records in this book. 

Another problem faced in this study is with regards to records of species from the Taiwan Straits. 

Unless the paper makes it clear from which part of the straits the specimens were collected from (i. e. 

if it is within Taiwanese waters), we have opted to exclude these records. In a few cases (e. g. some 

records in Fang, 1991), the precise collection point is stated and a clear decision can be made. In other 

cases (e. g. Huang, 1994 ), no precise information is provided, si& we have chosen to not to include 

them under Taiwan's fauna. Nevertheless, these taxa are still listed (under the discussion for the 

family) so that the reader is aware of them. In cases when the species is already known from Taiwan via 

another study; we have, for convenience, included references like Huang (1994) in the main list. 

The term "aut. " is utilised when an incorrect name had been used earlier but the taxa concerned are 

not synonyms. When names are subjective or objective synonyms, the authorship and year of the junior 

synonym is provided. In this instance, we have only included junior synonyms which have been used 

widely or for the Taiwanese fauna in the past; the synonymy of some species being far too long to list 

exhaustively. When no indication is provided for the species in question, it means that the name has 

been used in other common taxonomic combinations in the past, or that the previous spelling used by 

others are incorrect. All species citations in the text are cited in full and arranged in chronological and 

alphabetical order respectively. The nominal subfamily for each family is always listed first, with the 

rest arranged in alphabetical order. Within each subfamily, the genera and species are listed in alphabetical 

order. When species are recorded from Taiwan for the first time, additional information on the 

specimens used is listed in Appendix 1. For the convenience of Taiwanese carcinologists, we have provided 

Chinese translations for key localities (Appendix 2); and the original Chinesejjournal titles~which 

we had translated into English in the references (Appendix 3). We have also provided photographs of all 

new records, species whose live colours are not yet known or for taxa which we feel are interesting. 

Some of the records are based on old specimens whose original colours have faded and to show the readers 

what the fresh colours are, we have used photographs of specimens collected from other areas. 

The following institutional abbreviations are used-TMCD: National Taiwan Museum, Taipei; 

NTOU: National Taiwan Ocean University, Keelung; NKMT: National Kaohsiung Institute of Marine 

Technology, Kaohsiung; and ZRC: Zoological Reference Collection of the Raffles Museum, Singapore. 

General notes on classification used 

The family classification essentially follows that proposed by Ng (1998), with several amendments. 

The families Dairidae Ng and Rodriguez, 1986, and Matutidae De Haan, 1835, are recognised as separate 

families (Ng and Rodriguez, 1986; Bellwood, 1996). Since Ng (1998), several other families have 

also been recognised, although none are known from Taiwan thus far. Bellwood (1996) also recognises


the Hepatidae Stimpson, 1871, as a good family, which is reasonable, considering the comments of 

Guinot (1977, 1978) about Hepatus and its allies. For the Raninidae, Goeke (1981) had recognised a 

third subfamily, Symethinae, but Tucker (1998) argued that it should be regarded as a distinct family, 

the Symethidae Goeke, 1981. We prefer instead, however, to follow the classification of Guinot 

(1993 ) who recognised more subfamilies of Raninidae and regarded the Symethinae only as a subfamily. 

In a reappraisal of some unusual genera (e. g. Phyllotymolinum and Genkaia) previously placed in 

the Cyclodorippidae, Tavares (1998 ) established that they were sufficiently different to warrant the 

recognition of a distinct family, the Phyllotymolinidae Tavares, 1998. Recent developments specific to 

each family or subfamily are discussed under the respective taxa. The family Eumedonidae Dana, 1852, 

is here regarded as a subfamily of the Pilumnidae Samouelle, 1819, following the reappraisal by Ng 

and Clark (2o6\). 

We have decided against using subgenera in this treatment as this taxonomic unit is not often poorly 

defined and its use is often far too subjective. While subgenenc names are nomenclaturally valid, we 

have omitted them for practical purposes. When there is sufficient basis to recognise these subgenera as 

distinct genera (e. g. in the Sesarminae), and we have followed the consensus. In many cases, however, 

subgenera are simply regarded as full genera for convenience. For some families, the use of subgenera 

is very common (especially in the Portunidae, Grapsidae and Ocypodidae), but in most cases, the 

subgeneric limits are not well defined, and their use is difficult. As such, we have preferred not to use 

these subgeneric names for the moment. The same philosophy is followed for subspecies (see discussion 

for Uca, Ocypodidae). 

For the authorities and years for most of the suprageneric taxa, we mainly follow Manning and 

Holthuis (1981) and Ng (1998). The date of a key paper by Rathbun (1931), often incorrectly cited as 

" 1929", has been discussed in depth by Ng and Liu (1999). We also follow Ng and Ahyong (2001) in 

literature pertaining to MacLeay. There have been some confusion with the correct authorships and dates 

of publication of many of the species described by Jacquinot, Adams and White. The authors and dates 

used here are based on the historical work done by P. Clark (personal communication) (see also Clark 

and Presswell, 2001). 

Checklist 

Family Dromiidae De Haan, 1833 

Conchoecetes arttficiosus (Fabricius, 1798)—Chang, 1963; Ng, Chan and Wang, 2000 

Conchoecetes intermedius Lewinsohn, 1984 — Ng, Chan and Wang, 2000 

Cryptodromia tumida Stimpson, 1858 (= Cryptodromia fallax aut, )—Ng, Chan and Wang, 2000; 

McLay, Chan and Jeng, in press [This species was reported as C. fallax (Lamarck, 1818) by 

Ng, Chan and Wang (2000) but a recent reappraisal indicates that it should be referred to C. 

tumida instead. ] 

Cryptodromia fukuii (Sakai, 1936) (~ Peialomera fukuii) —Jeng, Jan, Tzang, Feng and Yang, 

1994, 1996; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 

1997; Jeng, Shao, Tzeng, Feng and Wu, 1998 

Dromia dormia (Linnaeus, 1763)—Ng, 1998; Ng, Chan and Wang, 2000 

Dromia wilsoni (Fulton and Grant, 1902) — Ng, Chan and Wang, 2000


Lauridromia dehaani (Rathbun, 1923) (= Dromia dehaani)—Chang, 1963; Sakai, 1976; Dai, 

Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994; Ng, 1998; Ng, Chan and 

Wang, 2000 

Lauridromia intermedia (Laurie, 1906)—Lin, 1949; Huang, 1994; Ng, Chan and Wang, 2000 

Petalomera granulata Stimpson, 1858—Ng, Chan and Wang, 2000; McLay, Chan and Jeng, in press 

Sphaerodromia ducoussoi McLay, 1991 — McLay, Chan and Jeng, in press 

Sphaerodromia kendalli (Alcock and Anderson, 1894)—Jeng, 1997; Ng, Chan and Wang, 2000 

Takedromia cristatipes (Sakai, 1969)—Ng, Chan and Wang, 2000 

Family Dynomenidae Ortmann, 1892 

Dynomene hispida Gueiin-Meneville, 1832 — Horikawa, 1940; Lin, 1949; Dai, Yang, Song and 

Chen, 1986; Dai and Yang, 1991; Huang, 1994; McLay, 1999 

Metadynomene tanensis (Yokoya, 1933 ) ( = Dynomene praedator aut. ) — Miyake, 1938; Lin, 

1949; McLay, 1999; McLay, Chan and Jeng, in press 

Family Homolidae De Haan, 1839 

Homola mieensis Sakai, 1979 — present record [Fig. la] 

Homola orientalis Henderson, 1888 — Jeng, Shao, Tzeng, Feng and Wu, 1998; Tan, Huang and 

Ng, 2000 

Homolochunia gadaletae Guinot and Richer de Forges, 1995 — Huang and Hsueh, 1998; Tan, Huang 

andNg, 2000 

Homolomannia occlusa Guinot and Richer de Forges, 1981 — Tan, Huang and Ng, 2000 

Homolomannia sibogae Ihle, 1912 — Tan, Huang and Ng, 2000 

Lqmoha murotoensis (Sakai, 1979)—Guinot and Richer de Forges, 1995; Tan, HuangandNg, 2000 

Latreillopsis bispinosa Henderson, 1888 — Ng and Huang, 1997; Tan, HuangandNg, 2000 

Moloha majora (Kubo, 1936)—Tan, HuangandNg, 2000 

Paromola japonica Parisi, 1915 — Jeng, Jan, Tzeng, Feng and Yang, 1994, 1996; Jeng, Jan, 

Tzeng and Feng, 1997; Ng and Huang, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng, 

1998; Tan, HuangandNg, 2000 

Paromola macrochira Sakai, 1961—Ng and Huang, 1997; Tan, HuangandNg, 2000 

Family Latreilliidae Stimpson, 1858 

Eplumula phalangium (De Haan, 1839) ( = Latreillia phalangium)—Tan, HuangandNg, 2000 

Latreillia valida De Haan, 1839 — Tan, Huang and Ng, 2000 

Family Raninidae De Haan, 1839 

Subfamily Ranininae De Haan, 1839 

Ranina ranina (Linnaeus, 1758 ) — Oshima, 1921b; Balss, 1922b; Maki and Tsuchiya, 1923; 

Sakai, 1937; Lin, 1949; Chang, 1965; Wang and Chen, 1981; Dai, Yang, Song and Chen, 

1986; Dai and Yang, 1991; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 1994; Ho, 1996; 

Jeng, Jan, Tzeng, Feng and Yang, 1996; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, 

1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng, 1998; Ng, Chan and Wang, 2000


Subfamily Lyreidinae Guinot, 1993 

Lyreidus stenops Wood-Mason, 1887 ( = Lyreiduspolitus Parisi, 1914) — Chang, 1963; Ng, Chan 

and Wang, 2000 

Lyreidus tridentatus De Haan, 1841 — Ng, Chan and Wang, 2000 

Subfamily Notopodinae Serene and Umali, 1972 

Cosmonotus grayi White, 1848 —Sakai, 1937; Sakai, 1976; Dai, Yang, Song and Chen, 1986; Dai 

and Yang, 1991; Ng, Chan and Wang, 2000 

Notopus dorsipes (Fabricius, 1798)—Ng, Chan and Wang, 2000 

Ranilia orientalis Sakai, 1963 — Ng, Chan and Wang, 2000 

Subfamily Raninoidinae Lbrenthey and Beurlen, 1929 

Notosceles serratifrons (Henderson, 1893) ( = Raninoides serratifrons) — present record [Fig. lc] 

[This species was originally described from the genus Raninoides, but is now clasified in Notosceles 

(see Guinot, 1993). ] 

Family Dorippidae MacLeay, 1838 

Subfamily DorippinaeMacLeay, 1838 

Dorippe quadridens (Fabricius, 1793) (= Dorippe dorsipes Miers, 1884, Dorippe frascone aut. 

partim) — Maki and Tsuchiya, 1923; Lin, 1949; Chang, 1963; Wang and Chen, 1981; Dai, 

Yang, Song and Chen, 1986; Holthuis and Manning, 1990; Dai and Yang, 1991; Huang, 1994; Lai, 

Huang and Fang, 1997; Chou, Lai and Fang, 1999 [Holthuis and Manning (1990: 18) include 

Lin's (1949) record of "D. dorsipes" under D. quadridens but in their discussion of D. sinica 

Chen, 1980, they list Lin's (1949) record from Taiwan under the distribution for this species. 

They did not discuss their reasons for placing Lin's record under one or the other species. Holthuis 

and Manning (1990: 29) comment that the distribution of the two species overlap in southern 

China (see discussion for D. sinica). We have examined specimens from Tahsi in northeastern 

Taiwan in the TMCD, NTOU and ZRC, and they are clearly D. quadridens as defined by Holthuis 

and Manning (1990) so it seems that the range of D. quadridens does extend into northern Taiwan 

and possibly into southern Japan as well. In fact, Holthuis and Manning (1990: 32) had included 

records of Maki and Tsuchiya (1923) from Keelung in northern Taiwan. Although D. sinica has 

yet to be formally reported from Taiwan, its presence there is almost certain, being present in 

Japan and southern China. As such, it would appear that the ranges of D. quadridens and D, sinica 

overlap over most of southern China and probably Taiwan. ] 

Dorippe sinica Chen, 1980 ( = Dorippe frascone aut. partim) — Holthuis and Manning, 1990 [Fig. 

Id] [ Dorippe sinica is a very distinct species and can easily be separated from D. quadridens by 

the characters enumerated by Holthuis and Manning (1990), and in the ZRC, there are specimens 

from southern China which are clearly referrable to this species. Holthuis and Manning included 

Lin's (1949) record of " D. dorsipes" under D. sinica as well without comment (see discussion 

for D. quadridens) and it is on the basis of this that D. sinica is included here for the Taiwan


fauna. It would not be surprising, however, as D. sinica has wide distribution in southern China 

and Japan (Chen, 1980; Holthuis and Manning, 1990). As such, we accept this record for the 

time being. ] 

Heikea arachnoides (Manning and Holthuis, 1986) — Ng and Huang, 1997 

Heikea japonicum (Von Siebold, 1824) ( = Dorippe japonica; Neodorippe {Neodorippe) japonica 

van taiwanensis Serene and Rohmimohtarto, 1969) — Maki and Tsuchiya, 1923; Lin, 1949; 

Chang, 1963; Serene and Rohmimohtarto, 1969; Sakai, 1976; Yang, 1979; Dai, Yang, Song 

and Chen, 1986; Holthuis and Manning, 1990; Dai and Yang, 1991; Lai, Huang and Fang, 

1997; Ng and Huang, 1997; Chou, Lai and Fang, 1999 •*• (fliyflWe, \1tt 

Paradorippe granulata (De Haan, 1841) (= Dorippe granulata) — Sakai, 1937, 1940; Lin, 1949; 

Chang, 1963; Takeda and Miyake, 1970; Sakai, 1976; Dai, Yang, Song and Chen, 1986; Holthuis 

and Manning, 1990; Dai and Yang, 1991; Huang, 1994 

Subfamily Ethusinae Guinot, 1977 

Fang (1991) reported Ethusa izuensis Sakai, 1937, from an unspecified location in the Taiwan 

Straits. Until this record can be substantiated, we prefer not to include it for Taiwan for the moment. 

Ethusa sexdentata (Stimpson, 1858)—- Ng and Huang, 1997 

Family Cyclodorippidae Ortmann, 1892 

Tymolus hirtipes Tan and Huang, 2000 — Tan and Huang, 2000 

Family Leucosiidae SamoueHe, 1819 

Fang ( 1991 ) reported numerous leucosiids from various unspecified locations in the Taiwan 

Straits. Of these, three have not been reported from Taiwan before, viz. Pseudophilyra woodmasoni 

Alcock, 1896, Nursilia tonsor Alcock, 1896, and Arcania globata Stimpson, 1858 (all Philyrinae). 

Huang (1994 ) also reported Arcania erinacea (Fabricius, 1787 ) from somewhere in the Taiwan 

Straits. Until these records can be substantiated, we prefer not to include them for Taiwan for the moment. 

Subfamily Leucosiinae Samouelle, 1819 

Leucosia anatum (Herbst, 1783) (= Leucosia longifrons De Haan, 1841) — Chang, 1963; Huang, 

1994; Jeng, Shao, Tzeng, Feng and Wu, 1998 [Fig. lh] ,^\ya\t, WUi 

Leucosia craniolaris (Linnaeus, 1758) — Lin, 1949; Sakai, 1976;/Dai, Yang, Song and Chen, 

1986; Dai and Yang, 1991; Huang, 1994; Jeng, Shao, Tzeng, Feng and Wu, 1998; Chou, Lai 

and Fang, 1999 

Leucosia formosensis Sakai, 1937 — Sakai, 1937; Horikawa, 1940; Lin, 1949; Chang, 1963; 

Sakai, 1976; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994 

Leucosia haematosticta Adams and White, 1849 — Huang, 1994; Jeng, Shao, Tzeng, Feng and Wu, 

1998 [Fig. le] 

Leucosia rhomboidalis De Haan, 1850 — Huang, 1994; present record [Fig. If] [Huang (1994) reported 

this species from the Taiwan Straits without indicating which side of the strait it was collected 

from. The present specimens are all from Taiwan proper. ]


Leucosia unidentata De Haan, 1841 — Chang, 1963; Huang, 1994; Jeng, Shao, Tzeng, Feng and 

Wu, 1998 

Leucosia vittata Stimpson, 1858 — Lin, 1949; Dai, Yang, Song and Chen, 1986; Dai and Yang, 

1991; Huang, 1994 [Fig. lg] 

Subfamily Ebaliinae Stimpson, 1871 

Heteronucia laminata (Doflein, 1904) — Ng and Huang, 1997 

Subfamily Philyrinae Rathbun, 1937 

Arcania elongata (Yokoya, 1933) — Ng and Huang, 1997 

Arcania heptacantha (De Haan, 1861 ) ( = Arcania siamensis Rathbun, 1909 ) — Lin, 1949 ; 

Chang, 1963; Wang and Chen, 1981; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; 

Huang, 1994 

Arcania quinquespinosa Alcock and Anderson, 1894 — Lin, 1949; Wang and Chen, 1981; Dai, 

Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994 

Arcania septemspinosa (Fabricius, 1787) — present record [Figs. 2a, b] 

Arcania undecimspinosa De Haan, 1841 (= Arcania granulosa Miers, 1877) — Sakai, 1937; Lin, 

1949; Chang, 1963; Sakai, 1976; Wang and Chen, 1981; Dai, Yang, Song and Chen, 1986; Dai 

andYang, 1991; Huang, 1994 

Iphiculus spongiosus Adams and White, 1849 — Lin, 1949 

Ixacylindrus (Fabricius, 1777) —Lin, 1949; Dai, Yang, SongandChen, 1986; Dai andYang, 1991; 

Huang, 1994 [Fig. 2c] 

Ixa edwardsii Lucas, 1858 — present record [Figs. 2d, e] [Dai, Yang, Song and Chen (1986) and 

Dai and Yang (1991) regard this species as synonymous with /. inermis but the available evidence 

suggests that both species should be recognised, at least for the time being (P. K. L. Ng and H. -L. 

Chen, unpublished data). ] 

Ixa inermis Leach, 1817 — Lin, 1949 

Ixoides cornutus MacGilchrist, 1905 — Lin, 1949; Wang and Chen, 1981; Dai, Yang, Song and 

Chen, 1986; Dai and Yang, 1991; Huang, 1994 [We recently obtained a female specimen of this 

species (TMCD) from Tungkang in southern Taiwan. In life, the animal is a uniform orange on all 

its dorsal surfaces. ] 

Myra afftnis Bell, 1855 — Chang, 1963 

Myra biconica Ihle, 1918 — present record [Fig. 2f ] 

Myrafugax (Fabricius, 1798) — Lin, 1949; Chang, 1963; Wang and Chen, 1981; Dai, Yang, Song 

and Chen, 1986; Dai and Yang, 1991; Huang, 1994; Jeng, Shao, Tzeng, Feng and Wu, 1998 

Nursiaplicata (Herbst, 1803) — present record [Fig. 2g] 

Parilia major Sakai, 1961 — Jeng, Shao, Tzeng, Feng and Wu, 1998; Tan, Wu and Huang, 2000 

Pariphiculus mariannae (Herklots, 1852) — present record [Fig. 2h] 

Philyra heterograna Ortmann, 1892 — present record [Fig. 3a] 

Philyrapisum De Haan, 1841 — Maki and Tsuchiya, 1923; Sakai, 1937, 1940; Lin, 1949; Chang, 

1963; Hsueh, 1996; Wang and Liu, 1996a, b; Ho and Hung, 1997; Jeng, Lin, Wang and Jan, 

1998; Lin, 1998; Wang and Liu, 1998; Jeng and Wang, 2000; Lee and Tung, 2000


Phityra platycheir De Haan, 1841 — Lin, 1949; Wang and Chen, 1981; Dai, Yang, Song and Chen, 

1986; Dai and Yang, 1991; Huang, 1994; Lai, Huang and Fang, 1997; Jeng, Shao, Tzeng, 

Feng and Wu, 1998; Chou, Lai and Fang, 1999 

Philyra tuberculosa Stimpson, 1858—Sakai, 1937, 1940; Lin, 1949; Chang, 1963; Sakai, 1976; 

Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994 

Randallia eburnea Alcock, 1896 — Chang, 1963 

Randallia pustuloides Sakai, 1961 — Jeng, Shao, Tzeng, Feng and Wu, 1998; Tan, Wu and Huang, 

2000 

Randallia pustulosa Wood - Mason and Alcock, 1891 — Tan, Wu and Huang, 2000 

Family Calappidae De Haan, 1833 

Calappa bicornis Miers, 1884 — present record [Fig. 3b] [See record of Calappa capellonis below. ] 

Calappa calappa (Linnaeus, 1758) — Jeng, 1997, 1998; Ng, 1998 [There are two well known 

"colour morphs" (one plain coloured, another spotted) [Figs. 3c, d] of this species (see Galil, 

1997), both of which occur in Taiwan. Morphological and genetic studies by P. K. L. Ng and 

T. -Y. Chan, however, have shown that these "morphs" are in fact distinct species, with the 

"spotted morph" belonging to a new taxon. This study is now in the final stages of completion. ] 

Calappa capellonis (Laurie, 1906) — Galil, 1997 [Figs. 3e, f] [Wang and Chen (1981) had listed 

Calappa bicornis from Taiwan; we have re-examined the specimen on which this record was based 

(a male (TMCD 212) from Taitung collected by C. -Y. Wei on 21 February 1961). The specimen 

in question is C. capellonis. In addition, we have also examined a recently collected male specimen 

(ZRC 2000. 2393) of C. capellonis collected from Tahsi on 10 April 2000. Calappa bicornis, 

however, is also present in Taiwan on the basis of present records. ] 

Calappa gallus (Herbst, 1803) —Lin, 1949; Chang, 1963; Dai, Yang, Song and Chen, 1986; Dai 

and Yang, 1991; Huang, 1994; Jeng, 1997, 1998 

Calappa hepatica (Linnaeus, 1758 ) ( = Cancer tuberculata Herbst, 1785 ) — Terasaki, 1902; 

Balss, 1922b; Maki and Tsuchiya, 1923; Sakai, 1937; Lin, 1949; Chang, 1963; Wang and 

Chen, 1981; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994; Wang and 

Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Galil, 1997; Jeng, 1997, 1998; Ng, 1998; 

Wang and Liu, 1998; Jeng, 2000 

Calappa japonica Ortmann, 1892 ( = Calappa flammea aut., incorrectly spelled "C. frammed' sometimes)— 

Horikawa, 1940; Ho, 1996; Galil, 1997 [This is the largest species of Calappa in Taiwan. 

Although C. calappa is supposed to grow to larger sizes than C. japonica, we have, as yet, 

not found such large specimens of C. calappa in Taiwan, In Taiwan, C. japonica seems to be 

more common in the southern part (e. g. Tungkang), regularly appearing in the markets for sale. ] 

Calappa lophos (Herbst, 1782)—Balss, 1922b; Lin, 1949; Chang, 1963; Wang and Chen, 1981; 

Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994; Ho, 1996; Galil, 1997; 

Ng, 1998; Chou, Lai and Fang, 1999; Ng, Chen and Chan, 1999 

Calappa philargius (Linnaeus, 1758) — Terasaki, 1902; Oshima, 1921a; Balss, 1922b; Maki and 

Tsuchiya, 1923; Wang and Chen, 1981; Dai, Yang, Song and Chen, 1986; Dai and Yang, 

1991; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 1994; Ho, 1996; Jeng, Jan, Tzeng, 

Feng and Yang, 1996; Galil, 1997; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Huang and


Liitzen, 1998; Jeng, Shao, Tzeng, FengandWu, 1998; Ng, 1998; Chou, LaiandFang, 1999 

Calappa pustulosa Alcock, "1896 — Galil, 1997; Jeng, Shao, Tzeng, FengandWu, 1998 

Calappa quadrimacuUtta Takeda and Shikatani, 1990 — Takeda and Shikatani, 1990; Jeng. 1997. 

1998; Huang and Liitzen, 1998; Ng, Chen and Chan, 1999 

Cycloes granulosa De Haan, 1837 (= Cryptosoma granulosa)—Lin, 1949; Dai, Yang, Song and chen, 

1986; Dai and Yang, 1991; Huang, 1994; Galil and Clark, 1996; Jeng, 1997, 1998; Jeng, 

Shao, Tzeng, FengandWu, 1998 

Mursia armata De Haan, 1837 — Jeng, Shao, Tzeng, FengandWu, 1998 [Fig. 3g] 

Mursia danigoi Galil, 1993 — Tan, Wu and Huang, 2000 

Mursia microspina Davie and Short, 1989 — Tan, Wu and Huang, 2000 

Paracyclois milneedwardsii Miers, 1886 — Tan, Wu and Huang, 2000 

Family Matutidae De Haan, 1833 

Ashtoret maculata (Miers, 1877) (= Matuta banksii aut., Matuta lunaris aut., part, Ashtoret lunaris 

aut. ) — Balss, 1922b; Maki and Tsuchiya, 1923; Sato, 1936d; Sakai, 1937; Lin, 1949; 

Chang, 1963; Wang, 1984; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994 

(part); Hsueh, 1996 (part); Jeng, Jan, Tzeng, Feng and Yang, 1996 (part); Jeng, 1997; 

Jeng, Jan, Tzeng and Feng, 1997 (part); Ng and Huang, 1997; Huang and Liitzen, 1998; Jeng, 

Shao, Tzeng, Feng and Wu, 1998; Lin, 1998 [The first author examined material sent to him by 

P. -W. Hsueh and this contains members of both A. maculata and M. victor] 

Izanami curtispina (Sakai, 1961) ( = Matuta curtispina aut. ) — Ng and Huang, 1997; Jeng, Shao, 

Tzeng, FengandWu, 1998; Chou, LaiandFang, 1999 

Matutaplanipes¥abncms, 1798 —Lin, 1949; Chang, 1963; Chang, 1965; Jeng, 1997; Ng, 1998 

Matuta victor (Fabricius, 1781) ( = Matuta banksii Leach, 1817, Matuta lunaris aut. ?) — Oshima, 

1921a; Balss, 1922b; Maki and Tsuchiya, 1923; Sakai, 1937; Lin, 1949; Chang, 1963; Dai, 

Yang, Song and Chen, 1986; Dai and Yang, 1991; Chang and Chen, 1992; Tzeng and Chen, 

1992; Huang, 1994 (part); Galil and Clark, 1994; Ho, 1996; Hsueh, 1996 (part); Jeng, Jan, 

Tzeng, Feng and Yang, 1996 (part); Wang and Liu, 1996a, b; Ho and Hung, 1997; Jeng, 

1997; Jeng, Jan, Tzeng and Feng, 1997 (part); Lai, Huang and Fang, 1997; Ng and Huang, 

1997; Jeng, 1998; Jeng, Lin, Wang and Jan, 1998; Jeng, Shao, Feng, Tzeng and Wu, 1998; 

Ng, 1998; Wang and Liu, 1998; Chou, LaiandFang, 1999; Jeng and Wang, 2000; Lee and 

Tung, 2000 

Family Majidae Samouelle, 1819 

The subfamilial system of the Majidae remains unsettled. While some subfamilies seem discrete, 

most are not clearly defined and many have problematic genera. We nevertheless follow Griffin and 

Tranter's (1986) system for the time being for want of anything better. 

Griffin and Tranter (1986) records several species from the Formosa Straits ( = Taiwan Straits), 

but none are included here under the Taiwanese fauna. For five species, Achaeus lacertosus Stimpson, 

1857, Pugettia intermedia Sakai, 1938 (Epialtinae), Hyastenus convexus Miers, 1884, H. diacanthus 

(De Haan, 1839) and H. hilgendorfi De Man, 1887 (Pisinae), co-ordinates were provided and 

they were clearly collected from parts of the straits which are outside Taiwanese jurisdiction. The data


for two species, Pugettia incisa (De Haan, 1839) and P. marissinica Takeda and Miyake, 1972 (both 

Epialtinae), were not sufficiently specific, but were probably collected in areas outside Taiwan as defined 

here. As such, tii^ also excluded for the moment. The presence of all four species in Taiwan, 

however, can be expected as they are relatively widespread taxa Fang (1991) reported numerous majids 

from various unspecified locations in the Taiwan Straits. Of these, three have not been reported from 

Taiwan before, viz. Maja compressipes (Miers, 1879) ( = M. brevispinosus Dai, 1981, see Chen 

and Ng, 1999) (Majinae), Hyastenus kyusyuensis (Yokoya, 1933) (Pisinae) and Achaeus tuberculatus 

Miers, 1879 (Inachinae). Until frhese wooFdA can be substantiated, we prefer not to include these 

records for Taiwan for the moment. The same is true for the records of Doclea rissoni Leach, 1815 (as 

D. gracilipes Stimpson, 1857) (Pisinae), Maja miersii Walker, 1887 (Majinae), and Micippa thalia 

(Herbst, 1803) (Mithracinae) by Huang (1994). 

A note on the valid name for the majine genus Thacanophrys Griffin and Tranter, 1986, is necessary. 

Griffin and Tranter (1986) established Thacanophrys for many species previously placed in the 

genera Chlorinoides Haswell, 1880, or Acanthophrys A. Milne Edwards, 1865; designating Chlorinus 

aculeataH. Milne Edwards, 1834, as its type species. These authors, however, appear to have 

forgotten the name Prismatopus Ward, 1933 (type species Prismatopus albanyensis Ward, 1933, bj^ 

monotypy). Since Chlorinus aculeata H, Milne Edwards, 1834, and Prismatopus albanyensis Ward, 

1933, are currently regarded as congeneric, Prismatopus Ward, 1933, has clear priority over Thacanophrys 

Griffin and Tranter, 1986, and must be used instead. 

Subfamily Majinae Samouelle, 1819 

Cyclax suborbicularis (Stimpson, 1858)—Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997 

Maja japonica Rathbun, 1932 ( - Maja nipponensis Sakai, 1934) — Lin, 1949; Chang, 1963; Dai, 


Maja kominatoensis Kubo, 1936 — Chou, Lai and Fang, 1999 

Maja spinigera De Haan, 1839 — Sakai, 1938a; Sakai, 1976;/Dai, Yang, Song and Chen, 1986; 

Dai and Yang, 1991; Huang, 1994 

Prismatopus longispinus (DeHaan, 1839) (= Chlorinoides longispinosus, Thacanophrys longispinus) 

— Suzuki, 1985 

Schizophroida hilensis (Rathbun, 1906 ) ( = Schizophroida manazuruana Sakai, 1933 ) — Chang, 

1963 

Schizopkrys aspera (H. Milne Edwards, 1834)—Miyake, 1938; Chang, 1963; Hwang and Yu, 1980; 

Jeng, 1997 

Subfamily Epialtinae MacLeay, 1838 

Acanthonyx formosa Wu, Yu and Ng, 1999 ( = Pugettia sp. aut. ?) — Horikawa, 1940; Wu, Yu 

andNg, 1999 • 

Huenia heraldica (De Haan, 1837 ) ( = Huenia proteus (De Haan, 1839 ), following Hokhuis, 

1987)—Jeng, 1997 

Menaethius monoceros (Latreille, 1825) — Sakai, 1938a; Dai, Yang, Song and Chen, 1986; Dai 

and Yang, 1991; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 1994, 1996; Jeng, 1997; 

Jeng, Jan, Tzeng and Feng, 1997; Jeng, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998


Xenocarcinus tuberculatus White, 1847 — Jeng, 1994b; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 

1997, 1998 

Subfamily Inachinae MacLeay, 1838 

Camposcia retusa Latreille, 1892 — Chang, 1963; Hwang and Yu, 1980; Tzeng and Chen, 1992; 

Jeng, Jan, Tzeng, Feng and Yang, 1994, 1996; Wang and Liu, 1996a; Yu, Jeng, Chan, Ho 

and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, 1998; Jeng, Shao, Tzeng, 

Feng and Wu, 1998; Wang and Liu, 1998 

Cyrtomaia curviceros Bouvier, 1915 — present record [Fig. 3h] [Ng and Huang (1997) reported this 

species from the Tungsha Islands, and its preserce on the main island is not at all surprising. ] 

Cyrtomaia murrayi Miers, 1886 — Ng and Huang, 1997 

Dumeia taiwanicus (Loh and Wu, 1998) ( = Pamtymolus taiwanicus) — Loh and Wu, 1998; Loh 

andNg, 1999 

Macrocheira kaempferi (Temminck, 1836) — Huang, YuandTakeda, 1990 

Platymaia bartschi Rathbun, 1916 — present record [Fig. 4a] [This species was reported from the 

Tungsha Islands by Ng and Huang (1997) but this is its first record from the main islands. 1 

Platymaia remifera Rathbun, 1916 ( = Platymaia alcocki aut. ) — Ng and Huang, 1997 ; Jeng, 

Shao, Tzeng, Feng and Wu, 1998 

PleistacantkajoryxOrtmann, 1893—Jeng, Shao, Tzeng, FengandWu, 1998; Chou, LaiandFang, 1999 / £fr 

Pleistacantha sanctijohannis Miers, 1879 — present record [Fig. 4c] [We have also recently examined 

recent specimens of this species (ZRC, NTOU) collected from Tungkang in southern Taiwan 

from depths of 200-300 m (now being studied by S. -H. Wu. ) In life, the species is bright orange 

on all its dorsal surfaces. ] 

Subfamily Mithracinae MacLeay, 1838 

Cyclocoeloma tuberculata Miers, 1880 — Jeng, 1997, 1998 [Fig. 4d] [This species is typically 

associated with coral reefs, covering itself with small anemones in life. ] 

Tiarinia cornigera (Latreille, 1825) — Tzeng and Chen, 1992 

Tiarinia depressa Stimpson, 1857 (= ? Tiarinia gracilis Dana, 1852, following Griifin and Tranter, 

1986) — Miyake, 1938; Chang, 1963; Suzuki, 1985; Huang, 1994; Jeng, 1997 

Tiarinia spinigera Stimpson, 1857 — Chang, 1963; Jeng, 1997 

Subfamily Pisinae Dana, 1851 

Doclea canalifera Stimpson, 1857 — Lin, 1949; Huang, 1994; Jeng, Shao, Tzeng, FengandWu, 

1998; Chou, Lai and Fang, 1999 ffoyjft )


such, we follow his classification here. ] 

Hyastenus diacanthus (De Haan, 1839) —Lin, 1949; Wang and Chen, 1981; Huang, 1994 

Phalangipus fUiformis Rathbun, 1916 ( = ? Loxorhynchus sp. ) — Maki and Tsuchiya, 1923; Griffin, 

1973; Sakai, 1976; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994; 

Chou, Lai and Fang, 1999 

Phalangipushystrix (Miers, 1886) (= Naxia hystrix, Naxioideshystrix) — Sato, 1936c, d; Dai, Yang, 

Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994 

Phalangipus longipes (Linnaeus, 1758)—Lin, 1949; Griffin, 1973; Huang, 1994; Jeng, Shao, 

Tzeng, Feng and Wu, 1998 fliiyiVp,Wj3 

Tylocarcinus styx (Herbst, 1803) — Miyake, 1938;AHuang, 1994 

Family Parthenopidae MacLeay, 1838 

Within the Parthenopidae, there have been several proposals for subfamilies, none of which are 

well defined. Ng and Rodriguez (1986) followed up on Guinot's (1977, 1978) suggestions and recognised 

several groupings which can be regarded as equivalent to subfamilies. Notably, the Daldorfiinae 

Ng and Rodriguez, 1986, and Aethrinae Dana, 1851, standout. Guinot and Bouchard (1998) recently 

regarded the Aethridae as a distinct family by itself. We follow their action here. The genus Cryptopodia 

and its allies also stand out in many respects and they are clearly not members of the Aethrinae as 

some believe (e. g. Serene, 1968; Sakai, 1976). For the time being, we prefer to refer them to their 

own subfamily, Cryptopodinae Stimpson, 1871. As discussed earlier, one of Ng and Rodriguez's 

(1986 ) groups, the Dairinae, is here regarded as a distinct family (see also Guinot and Bouchard, 

1998). 

Subfamily Parthenopinae MacLeay, 1838 

Aulacolambrus curvispinus (Miers, 1879) ( = Aulacolambrus brevibrachiatus aut. ) — Tan, Huang 

and Ng, 1999a [The record of A. brevibrachiatus by Tan, Huang and Ng (1999a) should be corrected. 

The types of almost all Aulacolambrus species have been recently re-examined as part of a 

revision by S. H. Tan, and the Taiwanese record should be reidentified as A. curvispinus instead. 

A. brevibrachiatus (Shen, Dai and Chen, 1982) is actually a junior synonym of A. granulosus 

(Miers, 1879). ] 

Aulacolambrus koplonotus (Adams and White, 1849)—Tan, Huang and Ng, 1999a 

Garthambrus stellata (Rathbun, 1906) — Tan, Huang and Ng, 1999a 

Parthenope longimanus (Linnaeus, 1764) — Balss, 1922b; Sakai, 1938a, 1940; Sakai, 1976; Wang 

and Chen, 1981; Chou, Lai and Fang, 1999; Tan, Huang and Ng, 1999a 

Platylambrus echinatns (Herbst, 1790) { = Parthenope echinatus; Platylambrus tuberculosus Stimpson, 

1857) —Maki and Tsuchiya, 1923; Sakai, 1938a, 1940; Lin, 1949; Chang, 1963; 

Huang, 1994; Tan, Huang and Ng, 1999a 

Platylambrus validus De Haan, 1839 — Chang, 1963; Tan, Huang and Ng, 1999a 

Rhinolambrus contrarius (Herbst, 1804)—Tan, Huang and Ng, 1999a 

Rhinolambrus lamellifrons (Adams andWhite, 1848) ( = Parthenope lamellifrons)— Dai, Yang, Song 

and Chen, 1986; Dai and Yang, 1991; Huang, 1994; Tan, Huang and Ng, 1999a 

Rhinolambrus nummiferus (Rathbun, 1906)—Jeng, Shao, Tzeng, Feng and Wu, 1998 

Rhinolambruspelagicus (Ruppell, 1830) — Balss, 1922b; Sakai, 1938a


Subfamily Cryptopodiinae Stimpson, 1871 

Cryptopodia fomicata (Fabricius, 1781) —Lin, 1949; Wang and Chen, 1981; Dai, Yang, Song and 

Chen, 1986; Dai and Yang, 1991; Huang, 1994; Chiong and Ng, 1998; Chou, Lai and Fang, 

1999; Tan, Huang and Ng, 1999a 

Heterocrypta transitans Stimpson, 1858 — Huang, 1994; Tan, Huang and Ng, 1999a 

Subfamily Daldorfiinae Ng and Rodriguez, 1986 

DaMorfia horrida (Linnaeus, 1758) (= Parthenope horridus) •—Maki and Tsuchiya, 1923; Sakai, 

1938a; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994; Tan, Huang and Ng, 

1999a; McLay, Chan and Jeng, in press 

Subfamily Lambrachaeinae Stevcic, 1994 

Lambrachaeus ramifer Alcock, 1895 — McLay, Chan and Jeng, in press 

Family Aethridae Dana, 1851 

Aethra scruposa (Linnaeus, 1764) —jTan, Huang and Ng, 1999a; McLay, Chan and Jeng, in press 

Sakaila japonica (Sakai, 1963)—McLay, Chan and Jeng, in press 

Family Dairidae Ng and Rodriguez, 1986 

Dairaperlata (Herbst, 1790) —Sakai, 1939; Lin, 1949; Chang, 1963; Hwang and Yu, 1980; 

Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997, 1998; Wang and Liu, 

1998 

Family Hymenosomatidae MacLeay, 1838 

Elamena truncata (Stimpson, 1858) — Ng and Chuang, 1996; Ng, Chen and Fang, 1999 ,^ 1 

Halicarcinus coralicola (Rathbun, 1909) (= Rhynchoplax coralicola)—Sakai, 1938a, 1976; JNg, 

Chen and Fang, 1999; Ng and Jeng, 1999a 

Halicarcinus hondai (Takeda and Miyake, 1971) ( = Rhynchoplax hondai) — Suzuki, 1985; Ng, 

Chen and Fang, 1999; Ng and Jeng, 1999a 

Halicarcinus setirostris (Stimpson, 1858)— Ng, Chen and Fang, 1999; Ng and Jeng, 1999a 

Trigonoplax unguiformis (De Haan, 1839)—Jeng, 1997, 1998; Ng, Chen and Fang, 1999; Ng and 

Jeng, 1999a 

Family Corystidae Samouelle, 1819 

Jonas choprai Serene, 1971 — Serene, 1971; Moosa, 1973; Yang, 1979; Ng, Chan and Wang, 

2000 

Jonas distincta (De Haan, 1835 ) — Horikawa, 1940; Lin, 1949; Chang, 1963; Sakai, 1976; 

Huang, 1994; Ng, Chan and Wang, 2000 

Jonas formosae (Balss, 1922) (= Gomeza distincta formosae, Jonas distincta formosensis [sic] or 

Jonas formosus [sic])—Balss, 1922a, c; Horikawa, 1940; Sakai, 1940; Lin, 1949; Sakai, 

1976; Yang, 1979; Chen, 1998; Ng, Chan and Wang, 2000 [The name for this species has been 

mis-spelt in many papers since Balss (1922a, c) ]


Family Atelecylidae Ortmann, 1893 

Trachycarcinus elegans Guinot and Sakai, 1970 — Ng and Chan, 1997 

Family Cancridae Latreille, 1803 

Chou, Lai and Fang (1999) reported Cancer amphioetus /from southwestern Taiwan, but we 

believe their record is most likely Cancer japonicus, which is the most common cancrid in Taiwanese 

waters. 

Cancer japonicus Ortmann, 1893 — Tan, Wu and Huang, 2000 

Cancer nadaensis Sakai, 1969 — Ng and Huang, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998 

Family Portunidae Rafinesque, 1815 

The subfamilial classification for the Portunidae follows that recently modified by Apel and Spiridonov 

( 1998 ) . Fang ( 1991 ) reported two species, Lupocyclus inaequalis (Walker, 1887 ) 

(Portuninae) and Charybdis vadorum Alcock, 1899 (Thalamitinae), from various unspecified locations 

in the Taiwan Straits. Until these records can be substantiated, we prefer not to include them for 

Taiwan for the moment. On the same rationale, we discount the records of Lupocyclus tugelae Barnard, 

1950 (Portuninae), Lissocarcinus polybioides Adams and White, 1849 (Caphyrinae), Liocarcinus 

corrugatus (Pennant, 1777) (as Macropipus corrugatus) (Polybiinae), and Charybdis vadorum Alcock, 

1899 (Thalamitinae), by Huang (1994) forthe time being. 

Subfamily Portuninae Rafinesque, 1815 

Lupocyclus philippinensis Semper, 1880 — Lin, 1949; Yu, 1979; Wang and Chen, 1981; Huang 

andYu, 1997 

Lupocyclus rotundatus Adams and White, 1849 — Lin, 1949; Yu, 1979; Huang and Yu, 1997; 

Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998 

Portunus argentatus (White, 1847) —Lin, 1949; Chang, 1963; Yu, 1979; Fang, 1991; Huang, 

1994; Huang and Yu, 1997; Jeng, Jan, Tzeng and Feng, 1997; Lai, Huang and Fang, 1997; 

Huang and Lutzen, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Chou, Lai and Fang, 1999 

Portunus gracilimanus (Stimpson, 1858) —Chang, 1963; Yu, 1979; Wang and Chen, 1981; Fang, 

1991; Huang, 1994; Huang and Yu, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, 

Tzeng, Feng and Wu, 1998; Chou, Lai and Fang, 1999 

Portunus granulatus (H. Milne Edwards, 1834) — Sakai, 1939; Chang, 1963; Dai, Yang, Song 

and Chen, 1986; Dai and Yang, 1991; Huang, 1994; Huang and Yu, 1997; Jeng, 1997,1998 

Portunus haanii Stimpson, 1858 ( - Portunus gladiator Fabricius, 1798) — Sato, 1936c, d; Lin, 

1949; Chang, 1963; Yu, 1979; Wang and Chen, 1981; Ho, 1996; Huang, 1994; Huang and 

Yu, 1997; Jeng, Jan, Tzeng and Feng, 1997; Huang and Lutzen, 1998; Jeng, Shao, Tzeng, 

Feng and Wu, 1998; Chou, Lai and Fang, 1999 

Portunus kastatoides Fabricius, 1798 — Maki and Tsuchiya, 1923; Sato, 1936c, d; Lin, 1949; 

Chang, 1963; Yu, 1979; Fang, 1991; Tzeng and Chen, 1992; Huang and Yu, 1997; Jeng, 

Jan, Tzeng and Feng, 1997; Lai, Huang and Fang, 1997; Jeng, Shao, Tzeng, Feng and Wu,


1998; Chou, Lai and Fang, 1999 

Portunus iranjae Crosnier, 1962 — Yu, Jeng, Chan, Ho and Shy, 1996; Huang and Yu, 1997; 

Jeng, 1997 

Portunus orbitosinus Rathbun, 1911 — Lin, 1949; Chang, 1963; Tzeng and Chen, 1992; Huang and 

Yu, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998 

Portunus pelagicus (Linnaeus, 1766) — Terasaki, 1903; Oshima, 1921a, b; Maki and Tsuchiya, 

1923; Sakai, 1939; Balss, 1922c; Miyake, 1938; Lin, 1949; Chang, 1963; Chang, 1965; Yu, 

1979; Wang and Chen, 1981; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 

1994; Jeng, Jan, Tzeng, Feng and Yang, 1994; Mo, 1996; Hsueh, 1996; Jeng, Jan, Tzeng, 

Feng and Yang, 1996; Wang and Liu, 1996a, b; Ho and Hung, 1997; Huang and Yu, 1997; 

Jeng, Jan, Tzeng and Feng, 1997; Lai, Huang and Fang, 1997; Jeng, 1998; Jeng, Shao, 

Tzeng, Feng and Wu, 1998; Ng, 1998; Wang and Liu, 1998; Chou, Lai and Fang, 1999 

Portunuspubescens (Dana, 1852) — Huang and Yu, 1997; Jeng, 1997 

Portunus pulchricristatus Gordon, 1931 — Huang and Yu, 1997 

Portunus sanguinolentus (Herbst, 1783) — Terasaki, 1903; Oshima, 1921a; Maki and Tsuchiya, 

1923; Sato, 1936c, d; Lin, 1949; Chang, 1963; Chang, 1965; Yu, 1979; Wang and Chen, 

1981; Wang, 1984; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994; 

Jeng, Jan, Tzeng, Feng and Yang, 1994; Ho, 1996; Jeng, Jan, Tzeng, Feng and Yang, 1996; 

Wang and Liu, 1996a, b; Yu, Jeng, Chan, Ho and Shy, 1996; Ho and Hung, 1997; Huang and 

Yu, 1997; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Lai, Huang and Fang, 1997; Huang 

and Ltitzen, 1998; Jeng, 1998; Jeng, Lin, Wang and Jan, 1998; Jeng, Shao, Fung, Tzeng and 

Wu, 1998; Ng, 1998; Wang and Liu, 1998; Chou, Lai and Fang, 1999; Lee and Tung, 2000 

Portunus trituberculatus (Miers, 1876) — Maki and Tsuchiya, 1923; Sakai, 1940; Chang, 1965; 4 f|f|^q}p 

Sakai, 1976; Yu, 1979; Jeng, Jan, Tzeng, Feng and Yang, 1994; Ho, 1996; Jeng, Jan, 

Tzeng, Feng and Yang, 1996; Huang and Yu, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, 

Shao, Tzeng, Feng and Wu, 1998; Ng, 1998; Chou, Lai and Fang, 1999 [Chou, Lai and Fang 

(1999) recorded this species as " P. tuberculosus" which is clearly a typographical mistake. ] 

Scylla olivacea (Herbst, 1796) (= Scylla serrata ml part) — Jeng, Lin, Wang and Jan, 1998; Keenan, 

Davie and Mann, 1998; Ng, 1998; Jeng and Wang, 2000; Lee and Tung, 2000 [Figs. 5a, b] 

Scylla paramamosain Estampador, 1949 ( = Scylla serrata aut. part) — Oshima, 1921a, b; Jeng, 

Jan, Tzeng, Feng And Yang, 1994, 1996; Wang and Liu, 1996a, b; Huang and Yu, 1997; Ho 

and Hung, 1997; Jeng, 1997, 1998; Keenan, Davie and Mann, 1998; Ng, 1998; Wang and Liu, 

1998; Chou, Lai and Fang, 1999; Jeng, 2000; Lee and Tung, 2000 [Figs. 5c, d] 

Scylla serrata (Forskal, 1775) — Keenan, Davie and Mann, 1998; Ng, 1998; Lee and Tung, 2000 

[Figs. 5e, f] [Records supposedly of this species by Terasaki, 1903; Balss, 1922c; Maki and 

Tsuchiya, 1923; Sato, 1936d; Sakai, 1939; Lin, 1949; Chang, 1963; Chang, 1965; Yu, 

1979; Wang and Chen, 1981; Su and Lue, 1984; Dai, Yang, Song and Chen, 1986; Dai and 

Yang, 1991; Shih, Lue and Wang, 1991; Kuo, 1995; Ho, 1996; Hsueh, 1996; Lai, Huang and 

Fang, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Lin, 1998 cannot be accurately identified 

as either no figures were provided, or they were not clear enough to allow an accurate identification; 

although their samples are probably mixed species lots]


Scylla tranquebarica (Fabricius, 1798) ( = Scylla serrata aut. part) — Keenan, Davie and Mann, 

1998; Ng, 1998; Lee and Tung, 2000 [Figs. 5g, h] 

Subfamily Caphyrinae Paulson, 1875 

Caphyra rotundifrons (A. Milne Edwards, 1869) — Jeng, 1994a; Yu, Jeng, Chan, Ho and Shy, 

1996; Huang and Yu, 1997; Jeng, 1997, 1998 

Lissocarcinus arkati Kemp, 1923 — present record [Fig. 4e] 

Lissocarcinus laevis Miers, 1886 — Lin, 1949; Chang, 1963; Wang and Chen, 1981; Jeng, Jan, 

Tzeng and Feng, 1997; Huang and Yu, 1997; Huang and Liitzen, 1998; Jeng, Shao, Tzeng, 

Feng and Wu, 1998; Ng and Jeng, 1999b 

Lissocarcinus orbiculare Dana, 1852 — Ng and Jeng, 1999b 

Subfamily Carupinae Paulson, 1875 

Catoptrus nitidus A. Milne Edwards, 1870 — Chou, Lai and Fang, 1999 

Libystes nitidus A. Milne Edwards, 1867 — Balss, 1922c; Sakai, 1940; Huang and Yu, 1997; Lai, 

Huang and Fang, 1997 

Subfamily Podophthalminae Dana, 1851 

Podophthalmus nacreus Alcock, 1899 — Lin, 1949; Huang and Yu, 1997 

Podophthalmus vigil (Fabricius, 1798) — Terasaki, 1904; Balss, 1922c; Maki and Tsuchiya, 

,„ 1923; Lin, 1949; Yu, 1979; Wang and Chen, 1981; Dai, Yang, Song and Chen, 1986; Dai 

and Yang, 1991; Huang, 1994; Huang and Yu, 1997; Jeng, Jan, Tzeng and Feng, 1997; Lai, 

Huang and Fang, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng, 1998; Chou, Lai and 

Fang, 1999 

Subfamily Polybiinae Ortmann, 1893 

Ovalipes punctatus (De Haan, 1833)—Yu, 1979; Wang and Chen, 1981; Ho, 1996; Huang and 

Yu, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng, 

1998 

Parathranites orientalis Miers, 1886 — Huang and Yu, 1997 

Subfamily Thalamitinae Paulson, 1875 

Charybdis acuta (A. Milne Edwards, 1869 )— Ho, 1996; Hsueh, 1996; Huang and Yu, 1997; 

Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Chou, Lai and 

Fang, 1999 

Charybdis acutifrons (De Man, 1879) — Ho, 1996; Huang and Yu, 1997; Lai, Huang and Fang, 

1997 

Charybdis affinis Dana, 1852 — Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 

1994; Huang and Yu, 1997; Lai, Huang and Fang, 1997; Ng, 1998; Chou, Lai and Fang, 1999 

Charybdis amboinensis Leene, 1938 — Ho, 1996; Huang and Yu, 1997; Jeng, Shao, Tzeng, Feng 

andWu, 1998


Charybdis anisodon (De Haan 1850) — Oshima, 1921b; Balss, 1922c; Maki and Tsuchiya, 1923; 

Sakai, 1939, 1940; Lin, 1949; Chang, 1963; Yu, 1979; Dai, Yang, Song and Chen, 1986; 

Dai and Yang, 1991; Huang, 1994; Huang and Yu, 1997; Lai, Huang and Fang, 1997; Ng, 

1998; Chou, Lai and Fang, 1999; Lee and Tung, 2000 

Charybdis annulate (Fabricius, 1798) —Balss, 1922c; Sakai, 1940, 1976; Tzeng and Chen, 1992; 

Yu, Jeng, Chan, Ho and Shy, 1996; Huang and Yu, 1997; Jeng, 1997; Lai, Huang and Fang, 

1997; Ng, 1998; Lee and Tung, 2000 

Charybdis bimaculata (Miers, 1886) — Sakai, 1939; Yu, 1979; Dai, Yang, Song and Chen, 1986; j. fllj^be 

Dai and Yang, 1991; Fang, 1991; Huang, 1994; Huang and Yu, 1997; Lai, Huang and Fang, 

1997; Huang and Lutzen, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998 

Charybdis feriatus (Linnaeus, 1758 ) ( = Charybdis cruciata Herbst, 1794 ) — Sato, 1936c, d; 

Sakai, 1939; Chang, 1963; Chang, 1965; Yu, 1979; Wang and Chen, 1981; Dai, Yang, Song 

and Chen, 1986; Dai and Yang, 1991; Ho, 1996; Huang, 1994; Wang and Liu, 1996a, b; Ho 

and Hung, 1997; Huang and Yu, 1997; Jeng, Jan, Tzeng and Feng, 1997; Lai, Huang and 

Fang, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng, 1998; Wang and Liu, 1998; Chou, 

Lai and Fang, 1999; Lee and Tung, 2000 

Charybdis granulata De Haan, 1835 — Ho, 1996; Huang and Yu, 1997; Jeng, Jan, Tzeng and 

Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Chou, Lai and Fang, 1999 

Charybdis hellerii (A. Milne Edwards, 1867) — Ho, 1996; Huang, 1994; Huang and Yu, 1997; 

Jeng, Jan, Tzeng and Feng, 1997; Lai, Huang and Fang, 1997; Jeng, Shao, Tzeng, Feng and 

Wu, 1998; Chou, Lai and Fang, 1999; Lee and Tung, 2000 

Charybdis hongkongensis Shen, 1934 — Huang, 1994; Huang and Yu, 1997; Chou, Lai and Fang, 

1999 

Charybdis japonica (A. Milne Edwards, 1861) — Maki and Tsuchiya, 1923; Sakai, 1939; Lin, 

1949; Chang, 1963, 1965; Sakai, 1976; Yu, 1979; Dai, Yang, Song and Chen, 1986; Dai and 

Yang, 1991; Huang and Yu, 1997; Lai, Huang and Fang, 1997; Jeng, Lin, Wang and Jan, 

1998; Ng, 1998; Chou, Lai and Fang, 1999; Jeng and Wang, 2000 

Charybdis lucifera (Fabricius, 1798) —Balss, 1922c; Sakai, 1940; Dai, Yang, Song and Chen, 

1986; Dai and Yang, 1991; Huang, 1994; Huang and Yu, 1997; Lai, Huang and Fang, 1997; 

Chou, Lai and Fang, 1999; Lee and Tung, 2000 

Charybdis miles (De Haan, 1835) —Chang, 1963; Yu, 1979; Wang and Chen, 1981; Dai, Yang, +%«te, f 

Song and Chen, 1986; Dai and Yang, 1991; Ho, 1996; Huang, 1994; Huang and Yu, 1997; 


Charybdis natator (Herbst, 1794) — Balss, 1922c; Maki and Tsuchiya, 1923; Lin, 1949; Chang, 

1963; Chang, 1965; Sakai, 1976; Yu, 1979; Hwang and Yu, 1980; Wang and Chen, 1981; ^ ^ ^ 

Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Ho, 1996; Huang and Yu, 1997; 

Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Lai, Huang and Fang, 1997; Jeng, 1998; Jeng, 

Shao, Tzeng, Feng and Wu, 1998; Ng, 1998; Chou, Lai and Fang, 1999 

Charybdis obtusifrons Leene, 1936 — Huang and Yu, 1997 

Charybdis orientalis Dana, 1852 — Jeng, Jan, Tzeng, Feng and Yang, 1994; Ho, 1996; Jeng, Jan, 

Tzeng, Feng and Yang, 1996; Huang and Yu, 1997; Jeng, Jan, Tzeng and Feng, 1997; Lai, 

Huang and Fang, 1997; Huang and Lutzen, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998


Charybdis sagamiensis Parisi, 1916 ( = Charybdis riversandersoni aut. ) — Ho, 1996 ; Hsueh, 

1996; Huang and Yu, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and 

Wu, 1998 [Takeda and Kubodera (1998) recently reiterated the fact that although Stephenson 

(1972) and Stephenson and Rees (1968) recognised C. sagamiensis Parisi, 1916 (type locality 

Japan), and C. riversandersoni Alcock, 1899 (type locality Indian Ocean), as separate species, 

most authors followed Sakai (1976) in regarding both as synonyms. Stephenson and Rees (1968) 

listed 27 characters which distinguish them, and Takeda and Kubodera (1998) concurred in recognising 

both as separate species. We have compared a specimen of C. riversandersoni from the Andaman 

Sea (male, 71. 6 by 49. 5 mm, ZRC 1999. 149, off Phuket, coll. April 1999) and numerous 

specimens of C. sagamiensis from Taiwan, and while most of the characters enumerated by 

Stephenson and Rees (1968) are not very reliable by themselves, they are useful when utilised as a 

suite. In particular, we find that the degree of inflation of the branchial region (distinctly more 

swollen in C. riversandersoni) is a useful character. Another reliable character is the last anterolateral 

tooth, which is distinctly more triangular and acutely triangular in C, sagamiensis, with that 

of C. riversandersoni appearing more truncate. ] 

Charybdis spiniferum (Miers, 1884) — Miers, 1886; Sakai, 1939 [This species was described from 

Queensland, Australia, with Miers (1886: 190) commenting that is was also found in "... South 

Formosa (Coll. Brit. Mus. ). This species is perhaps a variety of Goniosoma variegatum\ Leene 

(1938: 45-46) states, however, that "I think these two species [ C. spiniferum and C. hellerii] 

are also identical (as De Man already suggested in 1895). There are, however, some minor differences 

according to Henderson and De Man: a) the posterior border of the propodus of the natatory 

leg is not denticulated, b) the wrist has only two spinules on the outer surface instead of three spinules" 

. Nevertheless, she tentatively synonymised it under C. hellerii. Sakai (1939: 400), recognising 

C. spiniferum as a doubtful species, incorrectly attributed the authorship of the species to Fabricius. 

The synonymy of C. spiniferum and C. hellerii was followed by Serene (1968) but C 

spiniferum was not mentioned at all by Stephenson (1972 ) in his synopsis. In view of the differences 

between C. spiniferum and C. hellerii (sensu Miers, 1884; Henderson, 1893; Leene, 

1938), it may be better to recognise both as separate taxa for the time being. ] 

Charybdis truncata (Fabricius, 1798)—Lin, 1949; Chang, 1963; Yu, 1979; Wang and Chen, 

1981; Huang, 1994; Huang and Yu, 1997; Jeng, Jan, Tzeng and Feng, 1997; Lai, Huang and 

Fang, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng, 1998; Chou, Lai and Fang, 1999 

Charybdis variegata (Fabricius, 1798)—Chang, 1963; Yu, 1979; Huang, 1994; Huang and Yu, 

1997; Lai, Huang and Fang, 1997; Huang and Liitzen, 1998; Chou, Lai and Fang, 1999 

Thalamita admete (Herbst, 1803) — Wang, 1984; Suzuki, 1985; Dai, Yang, Song and Chen, 

1986; Dai and Yang, 1991; Huang, 1994; Yu, Jeng, Chan, Ho and Shy, 1996; Huang and Yu, 

1997; Jeng, 1997 

Thalamita chaptali (Audouin, 1826) — Yu, 1979; Huang and Yu, 1997; Lai, Huang and Fang, 

1997; Jeng, 1997 

Thalamita crenata (Latreille, 1829) (= Thalamita prymna aut. ) — Oshima, 1921a; Balss, 1922c; 

Maki and Tsuchiya, 1923; Lin, 1949; Chang, 1963; Yu, 1979; Jeng, Jan, Tzeng, Feng and 

Yang, 1994; Ho, 1996; Jeng, Jan, Tzeng, Feng and Yang, 1996; Huang and Yu, 1997; Jeng, 

s 1997; Jeng, Jan, Tzeng and Feng, 1997; Lai, Huang and Fang, 1997; Jeng, Shao, Tzeng, 

^+%*h 1133


Feng and Wu, 1998; Ng, 1998; Chou, Lai and Fang, 1999; Jeng, 2000; Lee and Tung, 2000 

Thalamita danae Stimpson, 1858 ( = Thalamita stimpsoni A. Milne Edwards, 1861 ) — Chang, 

1963; Yu, 1979; Hwang and Yu, 1980; Wang and Chen, 1981; Suzuki, 1985; Jeng, Jan, 

Tzeng, Feng and Yang, 1994; Ho, 1996; Jeng, Jan, Tzeng, Feng and Yang, 1996; Yu, Jeng, 

Chan, Ho and Shy, 1996; Huang and Yu, 1997; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; 

Jeng, Shao, Tzeng, Feng and Wu, 1998 

Thalamita demani Nobili, 1905 — Yu, Jeng, Chan, Ho and Shy, 1996; Huang and Yu, 1997; Jeng, 

1997 

Thalamita edwardsi Borradaile, 1900 — Maki and Tsuchiya, 1923; Yu, Jeng, Chan, Ho and Shy, 

1996; Huang and Yu, 1997; Jeng, 1997 

Thalamita gloriensis Crosnier, 1962 — Yu, Jeng, Chan, Ho and Shy, 1996; Huang and Yu, 1997; 

Jeng, 1997; Lai, Huang and Fang, 1997 

Thalamita holthuisi Stephenson, 1975 — Yu, Jeng, Chan, Ho and Shy, 1996; Huang and Yu, 1997; 

Jeng, 1997, 1998 

Thalamita Integra Dana, 1852 — Huang and Yu, 1997; Lai, Huang and Fang, 1997 

Thalamita kagosimensis Sakai, 1939 — Lin, 1949; Huang and Yu, 1997; Jeng, Jan, Tzeng and 

Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998 

Thalamitapelsarti Montgomery, 1931 (= Thalamita prymna aut. )—Ho, 1996 [Figs. 4g, h] 

Thalamita picta Stimpson, 1858 — Balss, 1922c; Suzuki, 1985; Dai, Yang, Song and Chen, 1986; 

Dai and Yang, 1991; Huang, 1994; Yu, Jeng, Chan, Ho and Shy, 1996; Huang and Yu, 1997; 

Jeng, 1997; 

Thalamita poissonii (Audouin and Savigny, 1817) — Balss, 1922c; Sakai, 1939, 1976; Huang and 

Yu, 1997; Chou, Lai and Fang, 1999 

Thalamita prymna (Herbst, 1803) (= Thalamita crenata aut. ) — Sato, 1936d; Sakai, 1939; + flliytlc?, r: 

Hwang and Yu, 1980; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Tzeng and 

Chen, 1992; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 1994, 1996; Yu, Jeng, Chan, 

Ho and Shy, 1996; Huang and Yu, 1997; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, 


Thalamita simaU. Milne Edwards, 1834 — Maki and Tsuchiya, 1923; Lin, 1949; Yu, 1979; Dai, tflliy^ltti 

Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994; Huang and Yu, 1997; Lai, 

Huang and Fang, 1997; Chou, Lai and Fang, 1999 

Thalamita spinimana Dana, 1852 — Yu, Jeng, Chan, Ho and Shy, 1996; Huang and Yu, 1997; 

Jeng, 1997, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng, 1998 

Thalamita stephensoni Crosnier, 1962 — Yu, Jeng, Chan, Ho and Shy, 1996; Huang and Yu, 1997; 

Jeng, 1997 

Thalamita wakensis Edmondson, 1925 — Yu, Jeng, Chan, Ho and Shy, 1996; Huang and Yu, 

1997; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998 

Family Carpiliidae Ortmann, 1893 

The genus Liagore De Haan, 1835, is presently in the family Carpiliidae, but it should not be 

classified there. Its characters are much closer to the Xanthidae as currently defined and should be

22 National Taiwan Museum Special Publication Series, No. 11, 86 pp.. April 2001 

transferred there (P. K. L. Ng, D. Guinot and P. J. F. Davie, ongoing study). For the moment, 

however, we retain it in this family for convenience. 

CarpUiusconvexus (Forsko\f 1775) — Balss, 1922c; Chang, 1963; Hwang and Yu, 1980; Wang and 

Chen, 1981; Dai, Yang, Song and Chen,' 1986; Dai and Yang, 1991; Huang, 1994; Wang and 

Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997, 1998; Ng, 1998; Wang and Liu, 

1998 

CarpUius maculatus (Linnaeus, 1758) — Sakai, 1939; Horikawa, 1940; Hwang and Yu, 1980; 

Wang and Chen, 1981; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994; 

Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and 

Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng, 1998; Wang and Liu, 1998 

Hagore rubromaculata De Haan, 1835 — Lin, 1949; Chang, 1963; Yang, 1979; Wang and Chen, 

1981; Huang, 1994; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 

1998 

Family Eriphiidae MacLeay, 1838 

The subfamilial system of the Eriphiidae utilised here requires more comment. Crosnier (in 

Serene, 1984) recognised three subfamilies, Eriphiinae, Oziinae and Dacryopiluminae. The family 

Eriphiidae can in fact be better split using the structures of the carapace, antenna, orbit and male second 

pleopod (P. K. L. Ng and S. K. Koh, unpublished data). To this effect, the subfamily Menippinae 

should also be recognised. Under this revised system, the Eriphiinae (species with orbital margin totally 

closed; antennae positioned some distance from orbits and antennules, being inserted at external frontal 

margin; distal part of male second pleopod tapers gradually to tip; distal part of male second pleopod 

distinctly shorter than subdistal part; right cheliped without special gastropod cutting tooth) contains 

only the genus Eriphia (one aberrant species, Eriphia norfolcensis Grant and McCulloch, 1907, was 

referred by Davie and Ng (2000) to a new genus, Bountiana, in the Oziinae). The Oziinae (species 

with orbital margin not or just closed; antennae positioned near orbits and antennules; distal part of male 

second pleopod tapers gradually to tip; distal part of male second pleopod subequal or longer than 

subdistal part; right cheliped of most genera with special gastropod cutting tooth) has the genera Ozius, 

Epixanthus, Epixanthoides, Lydia, Baptozius and Globopilumnus, The Menippinae (species with orbital 

margin not closed; antennae positioned near orbits and antennules; distal part of male second pleopod 

suddenly becoming very slender along terminal section; distal part of male second pleopod distinctly 

longer than subdistal part; right cheliped without special gastropod cutting tooth) has the genera 

Menippe, Myomenippe, Sphaerozius, Hypothalassia and Pseudocarcinus. Finally, the Dacryopiluminae 

(species with eyes positioned at lateral edge of trapezoidal carapace; frontal and anterolateral 

margins entire; orbits orbital margin not closed; antennae positioned some distance from orbits and 

frontal margin; distal part of male second pleopod suddenly becoming very slender along terminal 

section; distal part of male second pleopod longer than subdistal part; right cheliped without special 

gastropod cutting tooth) contains only the peculiar genus Dacryopilumnus. 

A note on the nomenclature of the genus Globopilumnus Balss, 1933, is necessary. While this 

name is relatively well known, it is in fact not the oldest available one and must be replaced by 

Eupilumnus Kossmann, 1877. Kossmann (1877) first established Eupilumnus as a subgenus of Pilum-


nus, and listed P. actumnoides A. Milne Edwards, 1873, P. nitidus A. Milne Edwards, 1873, P. 

longipes A. Milne Edwards, 1873, P. fissifrons Stimpson, 1858, P. dilatipes Adams and White, 

1849, P. vauquelinii Audouin, 1826, and P. savignyi Heller, 1861, under this name. No type 

species was designated then. Most workers have since closely associated the name Eupilumnus with 

Pilumnus Leach, 1815, and even regard them as synonyms. Rathbun (1930), however, had already 

selected Pilumnus actumnoides A. Milne Edwards, 1873, as the type species and this action is perfectly 

valid. As the type species of Globopilumnus Balss, 1933, is Pilumnus globosus Dana, 1852 (by original 

designation); and Pilumnus globosus Dana, 1852, and Pilumnus actumnoides A. Milne Edwards, 

1873, are both regarded as congeneric; this effectively makes Pilumnus (Eupilumnus) Kossmann, 

1877, a senior synonym of Globopilumnus Balss, 1933. It is also useful to note that Eupilumnus Kossmann, 

1877, is also a senior homonym of Eupilumnus Kingsley, 1880 (type species Eupilumnus websteri 

Kingsley, 1880), the latter which is now regarded as a junior synonym of the trapeziid Domecia 

Eydoux and Souleyet, 1842 (type species Domecia hispida Eydoux and Souleyet, 1842). 

Subfamily Eriphiinae MacLeay, 1838 

Eriphia scabricula Dana, 1852 — Sakai, 1939; Horikawa, 1940; Lin, 1949; Chang, 1963; Hwang 

and Yu, 1980; Suzuki, 1985; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Tzeng 

and Chen, 1992; Huang, 1994; Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; 

Jeng, 1997, 1998; Wang and Liu, 1998 

Eriphia sebana (Shaw and Nodder, 1803) (= Eriphia laevimana Guerin-Meneville, 1829)—Maki 

and Tsuchiya, 1923; Horikawa, 1940; Lin, 1949; Tzeng and Chen, 1992; Wang and Liu, 

1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; 

Lai, Huang and Fang, 1997, 1998; Ng, 1998; Wang and Liu, 1998 + W\yd&,\Ht3 

Eriphia smithii MacLeay, 1838 ( = Eriphia laevimana smithii)—Balss, 1922c; Maki and Tsuchiya, 

1923; Lin, 1949; Chang, 1963; Chang and Chen, 1992; Jeng, Jan, Tzeng, Feng and Yang, 

1996; Wang and Liu, 1996a; Jeng, Jan, Tzeng and Feng, 1997; Lai, Huang and Fang, 1997; 

Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng, 1998; Wang and Liu, 1998 [S. K. Koh and P. 

K. L. Ng (unpublished data) have shown that the real E. smithii is restricted to the western Indian 

Ocean, and specimens of " E. smithii" from the eastern Indian Ocean, Southeast Asia and eastern 

Pacific, including Taiwan, actually belong to a new species which is now being described. Until 

this study is published, we retain the name E. smithii for the Taiwanese specimens for convenience. 

] 

Subfamily Dacryopilumninae Serene, 1984 

Dacryopilumnus rathbunae Balss, 1932 — Suzuki, 1985 

Subfamily Menippinae Ortmann, 1893 

Hypothalassia armata (De Haan, 1835) — Jeng, Jan, Tzeng and Feng, 1997; Ng, 1998; Ho, Yu 

and Ng, 2000; Koh and Ng, 2000 

Menippe rumphii (Fabricius, 1798) — Ho and Hung, 1997; Jeng, Jan, Tzeng and Feng, 1997; Lai, 

Huang and Fang, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng, 1998; Chou, Lai and 

Fang, 1999; Lee and Tung, 2000


Myomenippe hardwickii (Gray, 1831) ( = Menippe granulosa De Man, 1888) — Lai, Huang and 

Fang, 1997 

Spkaerozius ntiidus Stimpson, 1858 — Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng 

and Wu, 1998; Ho, Yu and Ng, 2000 

Subfamily Oziinae Dana, 1851 

Epixanthus corrosus A. Milne Edwards, 1873 — Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; 

Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ho, Yu and Ng, 

2000 %«kt,iitf; 

Epixanthus dentatus (White, 1848) —Balss, 1922c; Sakai, 1940, 1976; Ng, 1998 

Epixanthus frontalis (H. Milne Edwards, 1834) — Balss, 1922c; Maki and Tsuchiya, 1923; Lin, 

1949; Hwang and Yu, 1980; Wang and Chen, 1981; Dai, Yang, Song and Chen, 1986; Dai and 

Yang, 1991; Chang and Chen, 1992; Jeng, Jan, Tzeng, Feng and Yang, 1994, 1996; Wang 

and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 

1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Wang and Liu, 1998 

Eupilumnus actumnoides (A. MilneEdwards, 1873) (= Pilumnus actumnoides, Globopilumnus actumnoides) 

— Suzuki, 1985 

Eupilumnus globosus (Dana, 1852) (= Globopilumnus globosus) — Balss, 1922c; Sakai, 1939; 

Sakai, 1976; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Yu, Jeng, Chan, Ho and 

Shy, 1996; Jeng, 1997 

Lydia annulipes (H. MilneEdwards, 1834) —Balss, 1922c; Sakai, 1939; Horikawa, 1940; Hwang 

andYu, 1980; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Chang and Chen, 1992; 

Tzeng and Chen, 1992; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 1996; Wang and Liu, 

1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; 

Lai, Huang and Fang, 1997; Jeng, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Wang and 

Liu, 1998 + Ifoyoltt fn%5 

Oziusguttatus H. MilneEdwards, 1834 —Ng, 1998 

Ozius rugulosus Stimpson, 1858 — Parisi, 1916; Maki and Tsuchiya, 1923; Sato, 1936d; Miyake, 

1938; Lin, 1949; Chang, 1963; Wang, 1984; Suzuki, 1985; Dai, Yang, Song and Chen, 

1986; Dai and Yang, 1991; Chang and Chen, 1992; Huang, 1994; Jeng, Jan, Tzeng, Feng and 

Yang, 1994, 1996; Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; 

Jeng, Jan, Tzeng and Feng, 1997; Jeng, Lin, Wang and Jan, 1998; Jeng, Shao, Tzeng, Feng 

andWu, 1998; Wang and Liu, 1998 

Ozius tuberculosa H. MilneEdwards, 1834 — Suzuki, 1985; Yu, Jeng, Chan, Ho and Shy, 1996; 

Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Lai, Huang and Fang, 1997; Jeng, Shao, 

Tzeng, Feng and Wu, 1998; Ng, 1998 

Family Xanthidae Macleay, 1S38 

The record of Palapedia nitida (Stimpson, 1858) (as Kraussia nitida) (Kraussiinae) by Huang 

(1994) from the Taiwan Straits is not included as we are uncertain from which side of the straits it was 

obtained from. Considering its wide distribution, its presence in Taiwan proper can be expected.


Subfamily Xanthinae MacLeay, 1838 

Demania cultripes (Alcock, 1898) —Ho, 1994 [Fig. 6a] 

Demania intermedia Guinot, 1969 — Ho, 1994 [Fig. 6b] 

Demania japonica Guinot, 1977 ( = DemaniarotundataauL partim) — ChenandNg, 1999 [Fig. 6c] 

Demania mortenseni (Odhner, 1925) — Ho, Yu and Ng, 2000 

Demania reynaudi (H. Milne Edwards, 1834)—Lin, 1949; Ho, 1994 [Fig. 6d] 

Demania rotundata Serene, in Guinot, 1969 ( = Demania reynaudi, Demania cultripes aut. ) — 

Serene, in Guinot, 1969; Takeda and Miyake, 1969; Sakai, 1976; Guinot, 1977; Guinot, 

1979; Yang, 1979; Garth and Ng, 1985; Dai, Yang, Song and Chen, 1986; Dai and Yang, 

1991; Ho, 1994; Huang, 1994; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng 

andWu, 1998; ChenandNg, 1999 [Fig. 6e] 

Demania scaberrima (Walker, 1887) — Wang and Chen, 1981; Huang, 1994; Ho, 1994 [Fig. 6f] 

Demania toxica Garth, 1971 — Ho, 1994 [Fig. 6g] 

Euryxanthops orientalis (Sakai, 1939) — Ho, Yu and Ng, 2000 

Juxtaxanthias lividus (Lamarck, 1818) — Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Ho, 

YuandNg, 2000 [Fig. 6h] 

Juxtaxanthias tetraodon (Heller, 1861) — Lin, 1949 

Lachnopodus bidentatus (A. Milne Edwards, 1867) — Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 

1997; Ho, Yu and Ng, 2000 

Lachnopodus rodgersi Stimpson, 1858 — Wang and Liu, 1996a; Wang and Liu, 1998 

Lachnopodus subacutus (Stimpson, 1858 ) — Jeng, Jan, Tzeng and Feng, 1997 ; Jeng, Shao, 

Tzeng, Feng and Wu, 1998; Ho, YuandNg, 2000 

Lachnopodus tahitensis De Man, 1889 — Ho, Yu and Ng, 2000 

Leptodius exaratus (H. Milne Edwards, 1834) —Balss, 1922c; Sakai, 1939; Chang, 1963; Hwang 

andYu, 1980; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Chang and Chen, 1992; 

Tzeng and Chen, 1992; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 1994, 1996; Wang 

and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 


Leptodius gracilis (Dana, 1852 ) — Chang and Chen, 1992; Jeng, Jan, Tzeng, Feng and Yang, 

1996; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, 


Leptodius nudipes (Dana, 1852) — Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Ho, Yu and 

Ng, 2000 

Leptodius sanguineus (H. Milne Edwards, 1834) — Parisi, 1916; Oshima, 1921a; Balss, 1922c; 

Maki and Tsuchiya, 1923; Lin, 1949; Chang, 1963; Hwang and Yu, 1980; Suzuki, 1985; Dai, 

Yang, Song and Chen, 1986; Dai and Yang, 1991; Chang and Chen, 1992; Jeng, Jan, Tzeng, 

Feng and Yang, 1996; Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Ho and 

Hung, 1997; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, 1998; Jeng, Shao, Tzeng, Feng 

andWu, 1998; Wang and Liu, 1998; Lee and Tung, 2000 

Macromedaeus crassimanus (A. Milne Edwards, 1867) — Ho, YuandNg, 2000 

Macromedaeus demani (Odhner, 1925) — Ho, YuandNg, 2000


Macromedaeus distinguendus (De Haan, 1835) — Sato, 1936d; Lin, 1949; Yu, Jeng, Chan, Ho 

and Shy, 1996; Jeng, 1997 

Neoxantkias impressus (Lamarck, 1818) — Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Ho, Yu 

andNg, 2000 

Neoxanthias michelae Serene and Vadon, 1981 — Ho, Yu and Ng, 2000 

Neoxanthops lineatus (A. Milne Edwards, 1867) — Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, 

Tzeng, Feng and Wu, 1998; Ho, Yu andNg, 2000 

Neoxanthops quadrilobatus (Sakai, 1939)—Yang, 1979; Ho, Yu and Ng, 2000[Fig. 7a] 

Paraxanthias notatus (Dana, 1852)—Lin, 1949; Chang, 1963; Suzuki, 1985; Yu, Jeng, Chan, 

Ho and Shy, 1996; Jeng, 1997 

Xanthias elegans (Stimpson, 1858) (= Paraxanthias elegans) — Miyake, 1938; Sakai, 1939; 

Sakai, 1976; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994; Jeng, Jan, 

Tzeng, Feng and Yang, 1994, 1996; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, 

Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998 + flntyak/ Mh 

Xanthias lamarcki (H. Milne Edwards, 1834) — Horikawa, 1940; Suzuki, 1985; Dai, Yang, Song 

and Chen, 1986; Dai and Yang, 1991; Huang, 1994; Yu, Jeng, Chan, Ho and Shy, 1996; 

Jeng, 1997 

Subfamily Actaeinae Alcock, 1898 

Actaea bocki Odhner, 1925 ( = Actaea ruppelli aut. ) — Balss, 1922c; Sakai, 1939, 1940, 1976; 

Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994 

Actaea savignyi (H. Milne Edwards, 1834)—Lin, 1949 

Actaeodes hirsutissimus (Riippell, 1830)—Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Ho, 

Yu and Ng, 2000 

Actaeodes tomentosus (H. Milne Edwards, 1834) —Sakai, 1939; Horikawa, 1940; Chang, 1963; 

Sakai, 1976; Wang, 1984; Suzuki, 1985; Dai, Yang, Song and Chen, 1986; Dai and Yang, 

1991; Chang and Chen, 1992; Tzeng and Chen, 1992; Huang, 1994; Jeng, Jan, Tzeng, Feng 

and Yang, 1996; Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; 

Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Wang and Liu. 

1998 A ifoy«to,*H3j; 

Forestia depressa (White, 1848)—Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and 

Wu, 1998; Ho, Yu and Ng, 2000 

Forestia scabra (Odhner, 1925) — Ng and Huang, 1997 

Gaillardiellus orientalis (Odhner, 1925) — Sato, 1936d 

Novactaea pulchetta (A. Milne Edwards, 1865) (= Actaea pulchella) —Miyake, 1938; Sakai, 

1939, 1976; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991 

Paractaea rufopunctata (H. Milne Edwards, 1834) — Miyake, 1938; Chang, 1963; Jeng, 1997 

Psaumis cavipes (Dana, 1852) —Sakai, 1939; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997 

Pseudoliomera helleri (A. Milne Edwards, 1865)—Ho, Yu and Ng, 2000 

Pseudoliomera lata (Borradaile, 1902)—Yu, Jeng, Chan. Ho and Shy, 1996; Jeng, 1997; Ho, Yu 

andNg, 2000

Ng et al. : Checklist of crabs from Taiwan 

Pseudoliomera remota (Rathbun, 1907)—Lin, 1949 

Pseudoliomera speciosa (Dana, 1852) (= Actaea speciosa)—Huang, 1994; Jeng, 1994b; Yu, 

Jeng, Chan, Ho and Shy, 1996; Jeng, 1997, 1998 

Subfamily Chlorodiinae Alcock, 1898 

Chlorodiella crispipleopa Dai, Yang, Song and Chen, 1986 — Dai, Yang, Song and Chen, 1986; 

Dai and Yang, 1991; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Ho, YuandNg, 2000 

Chlorodiella cytherea (Dana, 1852)—Sakai, 1976; Suzuki, 1985 * f0iyalt^W?3; 

Chlorodiella laevissima (Dana, 1852)—Miyake, 1938; Suzuki, 1985; Dai, Yang, Song and Chen, 

1986; Dai and Yang, 1991; Huang, 1994; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; 


Chlorodiella nigra (Forsskal, 1775) — Horikawa, 1940; Chang, 1963; Dai, Yang, Song and Chen, 

1986; Dai and Yang, 1991; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 1994, 1996; Yu, 

Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, 

Feng and Wu, 1998 

Phymodius granulosus (De Man, 1888) — Suzuki, 1985; Yu, Jeng, Chan, Ho and Shy, 1996; 

Jeng, 1997 

Phymodius nitidus (Dana, 1852)—Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; 

Jeng, 1997; Wang and Liu, 1998 

Pilodius areolatus (H. Milne Edwards, 1834)—Jeng, Jan, Tzeng, Feng and Yang, 1994, 1996; 

Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ho, 


Pilodius granulatus Stimpson, 1858 — Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng 

and Wu, 1998; Ho, Yu and Ng, 2000 

Pilodius paumotensis Rathbun, 1907-—Suzuki, 1985 

Pilodiuspilumnoides (White, 1848) — Suzuki, 1985 

Pilodius pugil Dana, 1852 — Lin, 1949 

Subfamily Cymoinae Alcock, 1898 

Cymoandreossyi (Audouin, 1826)—Jeng, 1994b; Yu, Jeng, Chan, HoandShy, 1996; Jeng, 1997 

CymomelanodactylusDeHzan, 1833 — Lin, 1949; Yu, Jeng, Chan, HoandShy, 1996; Jeng, 1997 

Cymo quadnlobatus Miers, 1884 — Yu, Jeng, Chan, HoandShy, 1996; Jeng, 1997; Ho, YuandNg, 

2000 

Subfamily Etisinae Ortmann, 1893 

Etisus anaglyptus H. Milne Edwards, 1834 — Miyake, 1938; Lin, 1949; Jeng, Jan, Tzeng and 


Etisus bargibanti Crosnier, 1987 — Ho, Yu and Ng, 2000 

Etisus dentatus (Herbst, 1785) —Yu, Jeng, Chan, HoandShy, 1996; Jeng, 1997; Ng, 1998; Ho, 


Etisus electra (Herbst, 1801) —Sakai, 1939; Dai, Yang, Song and Chen, 1986; Dai and Yang, 

27


1991; Huang, 1994; Jeng, 1997 

Etisus laevimanus Randall, 1840 —Sato, 1936d; Jeng, 1997; Ng, 1998; Jeng, 2000 

Etisus odhneri Takeda, 1971 —Jeng, 1997; Ho, Yu and Ng, 2000 

Etisus rhynchophorus A. Milne Edwards, 1873 — Ho, Yu and Ng, 2000 

Etisus splendidus Rathbun, 1906 — Ng and Huang, 1997; Ng, 1998 

Subfamily Euxanthinae Alcock, 1898 

Euxanthus exsculptus (Herbst, 1790) — Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Ho, Yu 

andNg, 2000 

Hypocolpus abbotii (Rathbun, 1894)—Ho, Yu and Ng, 2000 

Paramedaeus noelensis (Ward, 1934) — Suzuki, 1985 

Subfamily Kraussiinae Ng, 1993 

Kraussia rugulosa (Krauss, 1843) —Sakai, 1939; Sakai, 1976; Dai, Yang, Song and Chen, 1986; 

Dai and Yang, 1991; Huang, 1994; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997 

Palapedia Integra (De Haan, 1835) —Jeng, 1997; Ho, Yu and Ng, 2000 

Subfamily Liomerinae Sakai, 1976 

Actites erythra (Lanchester, 1901) ( = Liomera erythra) — Sakai, 1939; Horikawa, 1940; Lin, 

1949; Sakai, 1976 [Fig. 7b] [Odhner (1925) reported this species from the Taiwan Straits, but 

his record is just outside Taiwanese waters so is discounted (see Material and methods). The specimen 

figured here is from Singapore, the type locality of the species. ] 

Liomerabella (Dana, 1852)—Suzuki, 1985; Jeng, 1997 

Liomera cinctimana (White, 1847)—Lin, 1949; Yu. Jeng, Chan, Ho and Shy, 1996; Jeng, 1997 

Liomera laevis (A. Milne Edwards, 1873)—Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Ho, 

Yu andNg, 2000 

Liomera margaritata (A. Milne Edwards, 1873) — Lin, 1949; Yu, Jeng, Chan, Ho and Shy, 19%; 

Jeng, 1997 

Liomera monticulosa (A. Milne Edwards, 1873) — Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 

1997; Ho, Yu and Ng, 2000 

Liomera rubra {A. Milne Edwards, 1865) — Horikawa, 1940 

Liomera rugata (H. Milne Edwards, 1834)—Ho, Yu andNg, 2000 

Liomera venosa (H. Milne Edwards, 1834)—Sato, 1936d 

Neoliomera insularis (Adams and White, 1849) — Lin, 1949; Chang, 1963 

Subfamily Polydectinae Dana, 1851 

Lybia caestifera (Alcock, 1898) —Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Ho, Yu and 

Ng, 2000 

Lybia tessellata (Latreille, 1812)—Jeng, 1994b; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997. 

1998

Ng et al.: Checklist of crabs from Taiwan 29 

Subfamily Zalasiinae Serene, 1968 

Banareia armata A. Milne Edwards, 1869 — Miyake, 1938 

Banareia odhneri Sakai, 1974 — Ho, YuandNg, 2000 

Banareiasubgtobosa (Stimpson, 1858) — Ho, YuandNg, 2000 

CalvactaeatumidaV/axd, 1933 — Lin, 1949 

Zalasius dromiaeformis (De Haan, 1841)—Chang, 1963 

Zalasius sakaii Balss, 1938 — Ng and Huang, 1997 

Subfamily Zosiminae Alcock, 1898 

Atergatis dilatatus De Haan, 1835 — Hwang and Yu, 1980 

Atergatis floridus (Linnaeus, 1767) (= Atergatis ocyroe (Herbst, 1901))—Maki and Tsuchiya, 

1923; Horikawa, 1940; Wang, 1984; Suzuki, 1985; Dai, Yang, Song and Chen, 1986; Dai 

and Yang, 1991; Tzeng and Chen, 1992; Huang, 1994; Wang and Liu, 1996a; Yu, Jeng, 

Chan, Ho and Shy, 1996; Jeng, 1997, 1998; Wang and Liu, 1998 

Atergatis integerrimus (Lamarck, 1801)—Balss, 1922c; Jeng, 2000 

Atergatis latissimus (H. Milne Edwards, 1834)—Wang and Liu, 1996a; Wang and Liu, 1998 

Atergatis reticulatus De Haan, 1835 — Lin, 1949 

Atergatis subdentatus De Haan, 1835 — Ho, Yu and Ng, 2000 

Atergatopsis amoyensis De Man, 1879 ( = Actaea amoyensis) — Dai, Yang, Song and Chen, 1986; 

Dai and Yang, 1991; Huang, 1994; Ho, YuandNg, 2000 

Atergatopsis germaini A. Milne Edwards, 1865 — Tsai, Hwang, Chai and Jeng, 1996 

Atergatopsis signatus (Adams and White, 1849)—Ng, 1998; Ho, YuandNg, 2000 

Lophozozytnus incisus (H. Milne Edwards, 1834) — Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 

1997; Ho, YuandNg, 2000 

Lophozozymus pictor (Fabricius, 1798)—NgandChia, 1997 

Paratergatis langimanus Sakai, 1965 — Jeng and Ng, 1998 

Platypodia granulosa (Ruppell, 1830)— Lin, 1949; Dai, Yang, Song and Chen, 1986; Dai and 

Yang, 1991; Huang, 1994 

Platypodia semigranosa (Heller, 1861)—Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Ho, Yu 

andNg, 2000 

Pulchratis reticulatus Ng and Huang, 1997 — Ng and Huang, 1997 

Zosimus aeneus (Linnaeus, 1758) — Sakai, 1939; Horikawa, 1940; Hwang and Yu, 1980; Wang, 

1984; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Tzeng and Chen, 1992; Huang, 

1994; Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, 

Tzeng and Feng, 1997; Jeng, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Wang and Liu, 

1998 

Zozymodescavipes (Dana, 1852)—Ho, YuandNg, 2000 

Family Pilumnidae Samouelle, 1819 

The concept of this family has changed substantially since Balss (1933, 1957), and is in fact very 

similar to the tentative system suggested by Guinot (1978). Ng and Clark (2000, 2001) have discussed 

this matter in some depth (see also discussion for Goneplacidae later).


Subfamily Pilumninae Samouelle, 1819 

Fang (1991) reported Actumnusforficigerus (Stimpson, 1858) (Pilumninae), from an unspecified 

location in the Taiwan Straits. Until this record can be substantiated, we prefer not to include it for 

Taiwan for the moment. The same is true of the records of Pilumnus orbitospinis Rathbun, 1911, Actumnus 

dorsipes (Stimpson, 1858), A. forficigerus (Stimpson, 1858) (all three Pilumninae), Typhlocarcinus 

nudus Stimpson, 1858, Xenophthalmodes moebii Richters, 1880 (both Rhizopinae), and 

Parapanope euagora De Man, 1895 (in an unspecified subfamily as yet), by Huang (1994). 

Actumnus squamosus (DeHaan, 1835)—Jeng, Jan, Tzeng and Feng, 1997; Ho, YuandNg, 2000 

Actumnus pg. — Chou, Lai and Fang, 1999 [The paper describing this new species from southern 

Taiwan, which is allied to A. setifer (De Haan, 1835), by P.-H. Ho, H.-P. Yu and P. K. L. 

Ng, is now in press (2001)]. 

Benthopanope indica (De Man, 1887)—Jeng, Jan, Tzeng and Feng, 1997; Ho, Yu and Ng, 2000 

[Although Heteropanope pearsei Rathbun, 1932, described from Japan, is generally regarded as a 

junior synonym, unpublished studies (P. K. L. Ng) indicate that both are distinct species] 

Glabropilumnus laevis (Dana, 1852) — Suzuki, 1985 

Gorgonariana sodalis (Alcock, 1898)—Huang, 1994 

LophopUumnus diiatipes (Adams and White, 1849)—Ho, YuandNg, 2000 

Pilumnopeus makiana (Rathbun, 1931) (= Heteropanope makiana, following Balss, 1933; Davie, 

1989b) — Rathbun, 1931; Sakai, 1939, 1940, 1976; Dai, Yang, Song and Chen, 1986; Dai and 

Yang, 1991; Huang, 1994 + %*!&, M3j 

Pilumnus acanthosoma Ng, 2000 — Ng, 2000c [This new species, which is allied to P. dofleini 

Balss, 1933, from Japan, is known from a good series of specimens from southern Taiwan and the 

South China Sea . ] 

Pilumnus longicornis Hilgendorf, 1878 —Ho, YuandNg, 2000 

Pilumnus minutus De Haan, 1835 — Suzuki, 1985; Huang, 1994 

Pilumnus murphyi Ng, 1988 — Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Fung, Tzeng and 

Wu, 1998; Ho, YuandNg, 2000 

Pilumnus purpureus A. Milne Edwards, 1873—Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, 

Feng and Wu, 1998; Ho, Yu and Ng, 2000 

Pilumnus scabrisculus Adams and White, 1849 — Chang, 1963; Suzuki, 1985 

Pilumnus vespertilio (Fabricius, 1793) — Chang, 1963; Hwang and Yu, 1980; Tzeng and Chen, 

1992; Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997, 1998; Wang 

and Liu, 1998 

Subfamily Eumedoninae Dana, 1852 

Echinoecus pentagonus (A. Milne Edward, 1879) — Chia, Castro and Ng, 1999; Ng and Jeng, 

1999b 

Harrovia albolineata Adams and White, 1849 ( = Harrovia elegans aut. ) — Lin, 1949; Wang and 

Chen, 1981; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 1994, 1996; Jeng, 1997; Jeng, 

Jan, Tzeng and Feng, 1997; Chia and Ng, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng

Ng el al. : Checklist of crabs from Taiwan 31 

and Jeng, 1999b 

Permanotus purpureus (Gordon, 1934) ( = Harrovia purpurea) — Chia and Ng, 1998; Ng and 

Jeng, 1999b 

Tiaramedon spinosum (Miers, 1879) ( = Ceratocarcinus spinosuum) — Hwang and Yu, 1980; Ng 

and Jeng, 1999b 

Subfamily Galeninae A. Milne Edwards, 1862 

Galene bispinosa (Herbst, 1783) (= Galene granulata Miers, 1884)—Sakai, 1939; Lin, 1947, 

1949; Sakai, 1976; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994; 

Jeng, Jan, Tzeng and Feng, 1997; Ng, 1998 [Lin (1947) reported Galene granulata as a recent 

fossil record from Taiwan, but this taxon was originally described as an extant species. Galene 

granulosa Miers, 1884, however, is clearly a junior synonym of G. bispinosa (Herbst, 1783). 

The first author has examined a large series of specimens of this species from various parts of the 

Indo-West Pacific and G. granulosa is nothing more than a juvenile of G. bispinosa. Small specimens 

tend to have more granular carapaces, becoming distinctly smoother with increasing size. As 

such, both names must be synonymised. ] 

Subfamily Halimedinae Alcock, 1898 

Halimede fragifer (De Haan, 1835)—Lin, 1949; Huang, 1994 [ Halimede tyche (Herbst, 1801) is 

often regarded as a junior synonym of M. fragifer, but Galil (2000) recently argued that they were 

two distinct species. ] 

Subfamily Rhizopinae Stimpson, 1858 

Pseudolitochira Integra (Miers, 1884) (= Heteropilumnus integer)—Jeng, 1997 

Typhlocarcinus villosus Stimpson, 1858 — Lin, 1949; Huang, 1994 

Subfamily Tanaochelinae Ng and Clark, 2000 

Tanaocheles bidentata (Nobili, 1906) (= Chlorodiella bidentata)— Suzuki, 1985; Ng and Clark, 

2000 

Family Trapeziidae Miers, 1886 

Subfamily Trapeziinae Miers, 1886 

Quadrella maculosa Alcock, 1898 — Shin and Mok, 1996 

Tetralia glaberrima (Herbst, 1790)—Miyake, 1938; Horikawa, 1940; Jeng, Jan, Tzeng, Feng and 

Yang, 1994; 1996; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and 


Trapezia areolata Dana, 1852 — Horikawa, 1940; Lin, 1949; Suzuki, 1985; Tzeng and Chen, 

1992; Jeng, Jan, Tzeng, Feng and Yang, 1994, 1996; Jeng, Jan, Tzeng and Feng, 1997; Jeng, 


Trapezia cheni Galil, 1983 — Galil, 1983; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997 

Trapezia cymodoce (Herbst, 1799) — Horikawa, 1940; Lin, 1949; Hwang and Yu, 1980; Suzuki, 

1985; Jeng, Jan, Tzeng, Feng and Yang, 1994, 1996; Yu, Jeng, Chan, Ho and Shy, 1996;


Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998 

Trapezia digitalis Latreille, 1825 — Horikawa, 1940; Lin, 1949; Suzuki, 1985; Yu, Jeng, Chan, Ho 

and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 

1998 

Trapezia formosa Smith, 1869 — Chang, Chen and Chen, 1987; Yu, Jeng, Chan, Ho and Shy, 1996; 

Jeng, 1997 

Trapezia garthi Galil, 1983 — Galil, 1983; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997 

Trapezia guttata Riippell, 1830 — Jeng, 1994b; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997, 

1998 

Trapezia lutea Castro, 1997 ( = Trapezia cymodoce aut. part) — Jeng, 1994b; Wang and Liu, 

1996a; Castro, 1997; Jeng, 1998; Wang and Liu, 1998 

Trapeziarufopunctata (Herbst, 1799)—Jeng, 1994b; Yu, Jeng, Chan, HoandShy, 1996; Jeng, 1997, 

1998 

Trapezia septata Dana, 1852 — Galil and Lewinsohn, 1985; Jeng, 1994b; Wang and Liu, 1996a; 

Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, 


Trapezia serenei Odinetz, 1984 ( = Trapezia ferruginea aut., Trapezia dentata aut. ) — Lin, 1949; 

Hwang and Yu, 1980; Suzuki, 1985; Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 

1996; Jeng, 1997; Wang and Liu, 1998 

Trapezia speciosa Dana, 1852 — Yu, Jeng, Chan, HoandShy, 1996; Jeng, 1997 

Trapezia tigrina Eydoux and Souleyet, 1842 — Galil and Lewinsohn, 1984; Chang, Chen and Chen, 

1987; Jeng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998 

Subfamily Domeciinae Ortmann, 1893 

Domecia glabra Alcock, 1899 — Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997 

Domecia hispida Eydoux and Souleyet, 1842 — Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997 

Family Goneplacidae Macleay, 1838 

The subfamilial classification of what is now called the Goneplacidae is a major problem as it is 

clearly a polyphyletic grouping. Guinot (1971) was the last to appraise its composition but much 

remains unresolved. Ng ( 1987 ) formally removed the Rhizopinae s. str. into the Pilumnidae. Ng 

( 1987 ) rearranged the Rhizopinae and removed many genera from this subfamily which are not 

pilumnids. He tentatively retained these in the Goneplacidae s. lato. In this paper, for convenience, we 

refer the non-pilumnid genera previously placed in Rhizopinae to the Chasmocarcininae Serene, 1964. 

We otherwise follow Guinot's (1971) approximate subfamilial system for most of the Goneplacidae. 

We, however, concur with Richer de Forges (1996) who suggested that the genus Platypilumnus is 

closer to the Carcinoplacinae (Goneplacidae) than the Geryonidae (following Guinot, 1971). Similarly, 

Guinot (1971, 1978) regards the genera Halimede and Galene as pilumnids, an action we agree, 

and to this effect, we refer them to their respective subfamilies within the Pilumnidae. These 

subfamily groupings had in fact long been recognised (e. g. see Alcock, 1898, 1900), and there 

is good evidence that they are monophyletic. In addition, a new pilumnid subfamily (for Tanaocheles) 

has recently been described (Ng and Clark, 2000). Serene (1984) had proposed a classification of


the Pilumnidae which recognises five subfamilies: Pilumninae, Heteropanopeinae, Halimedinae, and 

for two new subfamilies, Planopilumninae and Heteropilumninae. Ongoing studies by the first author 

indicate that most of this classification is untenable. For example, there are no clear characters delineating 

the Pilumninae, Heteropanopeinae and Heteropilumninae, the latter which contains members of the 

the Pilumninae and Rhizopinae as currently understood. Certainly, as it now stands, many members of 

the Pilumninae and Heteropanopeinae cannot be effectively distinguished from each other and both are 

probably synonyms. With regards to the Planopilumninae, the type species of the type genus, 

Planopilumnus spongiosus (Nobili, 1905), is actually not a pilumnid at all but closer to goneplacids 

like the Pseudoziinae (P. K. L. Ng, unpublished data) instead. The genus Planopilumnus as currently 

understood, is heterogeneous. Additionally, Ng and Clark (2001) regarded the echinoderm-associates of 

the family Eumedonidae Dana, 1852, only as a subfamily of the Pilumnidae. 

The records of Carcinoplax meridionaiis Rathbun, 1923, Beteroplax nitida Miers, 1879, Heteroplax 

nagasakiensis Sakai, 1934, Ommatocarcinus macgillivrayi White, 1851, and Eucrate crenata 

(De Haan, 1835) by Huang (1994) are not included for the time being as it was merely stated as from 

the Taiwan Straits, without making clear from which side of the straits it was collected from. Carcinoplax 

suruguensis Rathbun, 1932, was also reported by Huang (1994) from somewhere in the Taiwan 

Straits, but we have examined specimens (now with J. -F. Huang) from northern Taiwan, so we include 

this record in our list. Similarly, Fang's (1991) record of Eucrate solans from an unspecified location 

in the Taiwan Straits is established in the present study through the first confirmed report of this 

species from Taiwan proper. 

Subfamily Goneplacinae MacLeay, 1838 

Ommatocarcinus pulcher (Barnard, 1950) — Ng and Huang, 1997 

Subfamily Carcinoplacinae H. Milne Edwards, 1852 

Carcinoplax longimana (De Haan, 1835) — Lin, 1949; Wang and Chen, 1981; Huang, 1994; 

Jeng, Jan, Tzeng, Feng and Yang, 1994; Ho, 1996; Jeng, Jan, Tzeng, Feng and Yang, 1996; 

Jeng, Jan, Tzeng and Feng, 1997; Huang and Liitzen, 1998; Jeng, Shao, Tzeng, Feng and Wu, 

1998; Ng, 1998 

Carcinoplax indica Doflein, 1904 — Huang, 1994; present record [Fig. 7c] [This species was reported 

from an unspecified location in the Taiwan Strait, but the present specimens are the first 

confirmed presence of the taxon from Taiwan proper. ] 

Carcinoplax suruguensis Rathbun, 1932 — Huang, 1994 [Fig. 7d] [As noted above, Huang (1994) 

reported this species from an unspecified location in the Taiwan Straits, but we have seen specimens 

from northern Taiwan and can thus confidently include it for the Taiawanese fauna. The taxonomy 

of this and other goneplacids, is now being studied by J. -F. Huang and P. -W. Hsueh. ] 

Carcinoplax tomentosa Sakai, 1969 — present record [Fig. 7eJ 

MathUdeUa serrata (Sakai, 1974) — Ng and Chan, 2000 

Platypilumnus soelae Garth, 1987 — Ng and Chan, 1998

34 National Taiwan Museum Special Publication Series, No. 11, 86 pp.. April 2001 

Psopheticus tnsignis Alcock, 1900 — Sakai, 1976 

Subfamily Chasmocarcininae Serene, 1964 

Camatopsis rubida Alcock and Anderson, 1899 —Fang, 1991: Huang, 1994 

Scalopidia spinosipes Stimpson, 1858 —Fang, 1991; Huang. 1994 [Fig. 7f. ] [The specimen figured 

is from the Gulf of Thailand, where the species can be very common. The species is normally 

associated with very soft substrates in relatively shallow waters less than 100 m depth. Smaller 

specimens tend to have more yellow and orange on the chelipeds. ] 

Subfamily Euryplacinae Stimpson, 1871 

Eucrate alcocki Serene, in Serene and Lohavanijaya, 1973 ( = Eucrate formosensis Sakai, 1974) 

— Sakai, 1974, 1976; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991 [Sakai (1974: 

94 ) described E. formosensis on the basis of one 29. 5 by 24. 0 mm male from Kaohsiung. 

Sakai (1976: 536) subsequently synonymised E. formosensis with E. alcocki Serene, 1971, 

noting that *'.. . the author found that E. formosensis is, save a few points of slight differences, 

quite identical with E. alcocki. " For the colour, he noted that ". . . of E. formosensis is similar to 

alcocki, but the central large spot is lacking. " Serene (1971: 916) based Eucrate alcocki on 

one male 25 by 20 mm from Nhatrang Bay in Vietnam. In his discussion for the species, he 

writes "Selon Sakai (1939), Heteroplax Stimpson, 1858, est un genre different d' Eucrate de 

Haan, 1835, Le type de la nouvelle espece correspond aux specimens indentifies Eucrate 

crenata var. dentata par Alcock ( 1900 ) et Chhapar (1957 ) et par erreur indentifi^s avec 

1' espece Heteroplax dentata Stimpson. 1858, qui est differente. La nouvelle espece alcocki remplace 

done dans la cle" de Campbell (1969 ) la forme mentionnee sous le nom de Eucrate dentata 

(Stimpson, 1858). " However, no description, diagnosis or figure was provided. Under the 

current zoological code, Serene's ( 1971 ) name "Eucrate alcocki" must be regarded as a 

nomen nudum. Serene and Lohavanijaya ( 1973 ) subsequently discussed this species, with a 

detailed comparison, statement of characters used and provision of figures. As such, the correct 

author citation and date of publication for this species must be Eucrate alcocki Serene, in 

Serene and Lohavanijaya, 1973. The authors are uncertain if the synonymy of E. alcocki and 

E. formosensis is correct. This is the only species in the region in which the anterolateral 

margin has only three teeth (including external orbital tooth) with the last being spiniform; and 

according to Dai and Yang (1991: 403), E. maculata Yang and Sun, 1979 (type locality 

China) is also a synonym. Sakai (1976: pi. 192 fig. 2) depicts an animal covered with numerous 

small spots, like the present specimens [Fig. 7g]. Eucrate alcocki, on the other hand, has a 

large median red gastric spot with a smaller spot on each side (Serene and Lohavanijaya, 

1973 ) . In any case, the genus Eucrate needs to be revised. For the moment, we retain 

Sakai's (1976) synonymy of the two species for lack of more conclusive evidence. ] 

Eucrate Solaris Yang and Sun, 1979 — present record [Fig. 7h] 

Subfamily Pseudoziinae Alcock, 1898 

Pseudozius caystrus (Adams and White, 1849) — Hwang and Yu, 1980; Ng and Wang, 1994; 

Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Shao,

Tzeng, Feng and Wu, 1998; Wang and Liu, 1998 


Family Geryonidae Colosi, 1923 

Chaceon granulates (Sakai, 1978)—Ng, 1998; Ng, Chan and Ng, 1998; Ng and Manning, 1998 

[Fig. 8a] [This large species is usually beige-brown in colour but is often very dark to almost black 

in parts, possibly because of their foraging near hydrothermal vents. We have observed and/or 

examined numerous specimens of various sizes from off northern Taiwan and Tiaoyutai Islands 

in the Peikuan Crab Museum; and smaller specimens may be maroon to almost white in 

colour. ] 

Chaceon manningi Ng, Lee and Yu, 1994 — Ng, Lee and Yu, 1994; Ng, Chan and Ng, 1998; 

Ng and Manning, 1998 [The first author has examined a dried mounted specimen of this ^~ ^ s 

species supposedly collected from southern Taiwan in the home of a fishermen in the port of ^ '"^ 

Tungkang near Kaohsiung. There is also at least one large dried male specimen of C. manningi CZ°°, 

obtained from off Tahsi in the Peikuan Crab Museum. Its presence in Taiwan, however, is not 

unexpected as the species was originally described from the Tungsha Islands in the South China 

Sea and is known from southern Japan as well (Ng and Manning, 1998). ] 

Family Ocypodidae Rafinesque, 1815 

With regards to the crabs of the genus Uca in this family, the standard reference remains Crane's 

(1975 ) masterpiece. Unfortunately, a good number of her new subgeneric, species and subspecies 

names are synonymous with those proposed by Bott (1973a, b). Bott's descriptions and discussions, 

however, are very brief and present a number of taxonomic problems. Because Crane's book was in 

press for many years, Bott's taxa were not included (or perhaps were not known to her) at the time her 

book went to press. Von Hagen (1976), in his review of Crane's book, commented that because of this 

confusion, it might be better just to use the broad genus name Uca for convenience. This has been followed 

by most workers since. Despite the nomenclatural and taxonomic problems associated with Crane's 

and Bott's taxa, there has been no concerted attempt to resolve them as yet. In any case, most workers 

have tended to follow Crane's scheme (and names) for these crabs, with Bott's works usually ignored. 

A second aspect of Uca taxonomy needs a brief comment, and that is with regards to Crane's concept of 

subspecies, which differs from modern ideas of what these taxa are. Nevertheless, we prefer not to use 

the subspecies concept here in the present compilation, partially for convenience, and mainly because 

even modern definitions of subspecies are still a matter of some debate. 

A note on the nomenclature of the macrophthalmine subgenus Macrophthalmus (Mopsocarcinus) 

Barnes, 1967, is necessary. Barnes(1967), in naming a new subgenus, Macrophthalmus(Mopsocarcinus) 

(type species Macrophthalmus boscii Audouin, 1826), was apparently unaware that there was an earlier 

name, Chaenostoma Stimpson, 1858 (type species Chaenostoma orientale Stimpson, 1858). Since 

Chaenostoma orientale Stimpson, 1858, is now regarded as a junior synonym of Macrophthalmus 

boscii Audouin, 1826, the name Chaenostoma Stimpson, 1858, must have priority. Although the 

name Chaenostoma has been used several times UA the use by Stimpson (1858) is the first we know of. ^ 

Subfamily Ocypodinae Rafinesque, 1815 

Ocypode ceratophthalmus (Pallas, 1772) — Parisi, 1918; Oshima, 1921a; Balss, 1922c; Maki and


Tsuchiya, 1923; Takahasi, 1934a, b, 1935; Sakai, 1939; Horikawa, 1940; Dai, Yang, Song 

and Chen, 1986; Fukui, Wada and Wang, 1989; Dai and Yang, 1991; Huang, Yu and Takeda, 

1992; Wu, 1992; Huang, 1994; Kuo, 1995; Hsueh, 1996; Wang and Liu, 1996a, b; Yu, 

Jeng, Chan, Ho and Shy, 1996; Ho and Hung, 1997; Jeng, 1997, 1998; Jeng, Lin, Wang and 

Jan, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Lin, 1998; Ng, 1998; Wang and Liu, 

1998; Jeng, 2000; Jeng and Wang, 2000; Lee and Tung, 2000 

Ocypode cordimana Desmarest, 1825 — Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; 

Huang, 1994 [Huang, Yang and Ng (1998) had shown that the Taiwanese records of " O. cordimana" 

they could check or assess all invariably belonged to O. sinensis instead. But since the real 

O. cordimana has a very wide Indo-West Pacific distribution, its presence in Taiwan is to be expected. 

The records of O. cordimana by Dai, Yang, Song and Chen (1986) and Dai and Yang 

(1991) from Taiwan are significant as some of these authors were the ones who described O. 

sinensis originally, and in their books, also clearly differentiate between the two species. In any 

case, we have on hand, one male specimen (33. 4 by 31. 5 mm, coll. 2 September 2000, 

TMCD) recently obtained from Renting in southern Taiwan which confirms that O. cordimana s. 

str. is actually present in Taiwan. ] 

Ocypode sinensis Dai, Song and Yang, 1985 ( = Ocypode cordimana aut., Ocypode stimpsoni aut. ) 

— Maki and Tsuchiya, 1923; Takahasi, 1934a, b, 1935; Sakai, 1939; Horikawa, 1940; Lin, 

1949; Chang, 1963; Sakai, 1976; Wang and Chen, 1981; Wang, 1984; Shih, Lue and Wang, 

1991; Chang and Chen, 1992; Huang, Yu and Takeda, 1992; Wu, 1992; Jeng, Jan, Tzeng, 

Feng and Yang, 1994, 1996; Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 

1997; Jeng, Jan, Tzeng and Feng, 1997; Huang, Yang and Ng, 1998; Jeng, 1998; Jeng, Shao, 

Tzeng, Feng and Wu, 1998; Lin, 1998; Ng, 1998 (part); Wang and Liu, 1998; Liu, 1999; 

Jeng, 2000; Lee and Tung, 2000 

Ocypode stimpsoni Ortmann, 1897 — Sakai, 1939, 1940; Chang, 1963; Sakai, 1976; Su and Lue, 

1984; Dai, Yang, Song and Chen, 1986; Fukui, Wada and Wang, 1989; Dai and Yang, 1991; 

' Shih, Lue and Wang, 1991; Chang and Chen, 1992; Huang, Yu and Takeda, 1992; Tzeng and 

Chen, 1992; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 1994; Kuo, 1995; Hsueh, 

1996; Jeng, Jan, Tzeng, Feng and Yang, 1996; Wang and Liu, 1996a, b; Ho and Hung, 1997; 

Jeng, Jan, Tzeng and Feng, 1997; Jeng, Lin, Wang and Jan, 1998; Jeng, Shao, Tzeng, Feng 

and Wu, 1998; Kitaura, Wada and Nishida, 1998; Lin, 1998; Wang and Liu, 1998; Jeng 

and Wang, 2000 

Uca arcuata (De Haan, 1835) — Parisi, 1918: Maki and Tsuchiya, 1923; Takahasi, 1934a, b, 

1935; Sato, 1936b, c; d; Sakai, 1939; Horikawa, 1940; Lin, 1949; Chang, 1963; Crane, 

1975; Sakai, 1976; Su and Lue, 1984; Wang, 1984; Dai, Yang, Song and Chen, 1986; Fukui, 

Wada and Wang, 1989; Huang, Yu and Takeda, 1989; Dai and Yang, 1991; Shih, Lue and 

Wang, 1991; Wu, 1992; Huang, 1994; Shih, 1994; Hsueh, 1995; Kuo, 1995; Hsueh, 1996; 

Wang and Liu, 1996a, b; Yu, Jeng, Chan, Ho and Shy, 1996; Ho and Hung, 1997; Jeng, 

1997, 1998; Jeng, Lin, Wang and Jan, 1998; Lin, 1998; Wang and Liu, 1998; Jeng and 

Wang, 2000; Lee and Tung, 2000; Shih, 2000a [The record of " U. formosensis" by Wu, Ling, 

Shieh and Wang (1962) from Tanshui is most likely V. arcuata. They describe this female specimen 

as having a very broad merus which is characteristic of U. arcuata and not U. formosensis.

Ng et at. : Checklist of crabs from Taiwan 37 

In any case, U. arcuata is the most common fiddler crab in Tanshui. ] 

Uca borealis Crane, 1975 ( = Uca vocans aut., Uca nttidus aut., Uca marionis aut., Uca vocans 

borealis) — Balss, 1922c; Maki and Tsuchiya, 1923; Takahasi, 1934a, b, 1935; Sakai, 1939; 

Crane, 1975; Sakai, 1976; Su and Lue, 1984; Dai, Yang, Song and Chen, 1986; Fukui, Wada 

and Wang, 1989; Huang, Yu and Takeda, 1989; Dai and Yang, 1991; Shih, Lue and Wang, 

1991; Tzeng and Chen, 1992; Wu, 1992; Huang, 1994; Shih, 1994; Kuo, 1995; Hsueh, 

1996; Wang and Liu, 1996a, b; Yu, Jeng, Chan, Ho and Shy, 1996; Ho and Hung, 1997; 

Jeng, 1997, 1998; Jeng, Lin, Wang and Jan, 1998; Lin, 1998; Wang and Liu, 1998; Jeng and 

Wang, 2000; Lee and Tung, 2000; Shih, 2000a 

Uca coarctata (H. Milne Edwards, 1852) — Fukui, Wada and Wang, 1989; Huang, Yu and Takeda, 

1989; Shih, 1994; Wang and Liu, 1996b; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 

1997, 1998; Lee and Tung, 2000; Shih, 2000a 

Uca crassipes (White, 1847 ) ( = Uca gaimardi H. Milne Edwards, 1852; Uca chlorophthalmm 

crassipes) — Sakai, 1939; Horikawa, 1940; Lin, 1949; Wang, 1984; Huang, Yu and Takeda, 

1989; Tzeng and Chen, 1992; Shih, 1994; Wang and Liu, 1996a, b; Yu, Jeng, Chan, Ho and 

Shy, 1996; Ho and Hung, 1997; Jeng, 1997, 1998; Wang and Liu, 1998; Jeng, 2000; Lee and 

Tung, 2000; Shih, 2000a 

Uca dussumieri (H. Milne Edwards, 1852 ) — Huang, Yu and Takeda, 1989; Tzeng and Chen, 

1992; Shih, 1994; Wang and Liu, 1996b; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997, 

1998; Lee and Tung, 2000; Shih, 2000a 

Uca formosensis Rathbun, 1921 — Rathbun, 1921; Maki and Tsuchiya, 1923; Takahasi, 1934a, b, 

1935; Sato, 1936c, d; Sakai, 1939; Horikawa, 1940; Sakai, 1940; Lin, 1949; Crane, 1975; 

Sakai, 1976; Su and Lue, 1984; Dai, Yang, Song and Chen, 1986; Fukui, Wada and Wang, 

1989; Huang, Yu and Takeda, 1989; Dai and Yang, 1991; Shih, Lue and Wang, 1991; Wu, 

1992; Huang, 1994; Shih, 1994; Kuo, 1995; Wang and Liu, 1996a, b; Ho and Hung, 1997; 

Shih, 1997; Jeng, Lin, Wang and Jan, 1998; Lin, 1998; Wang and Liu, 1998; Shih, Mok, 

Chang and Lee, 1999; Lee and Tung, 2000; Shih, 2000a 

Ucalactea (De Haan, 1835) — Parisi, 1918; Balss, 1922c; Maki and Tsuchiya, 1923; Takahasi, 

1934a, b, 1935; Sato, 1936a, c, d; Sakai, 1939; Horikawa, 1940; Lin, 1949; Chang, 1963; 

Crane, 1975; Sakai, 1976; Su and Lue, 1984; Wang, 1984; Dai, Yang, Song and Chen, 1986; 

Fukui, Wada and Wang, 1989; Huang, Yu and Takeda, 1989; Dai and Yang, 1991; Shih, Lue 

and Wang, 1991; Wu, 1992; Huang, 1994; Shih, 1994; Hsueh, 1995; Kuo, 1995; Hsueh 

1996; Wang and Liu, 1996a, b; Ho and Hung, 1997; Jeng, Lin, Wang and Jan, 1998; Jeng, 

Shao, Tzeng, Feng and Wu, 1998; Lin, 1998; Wang and Liu, 1998; Jeng, 2000; Jeng and 

Wang, 2000; Lee and Tung, 2000; Shih, 2000a 

Uca perplexa (H. Milne Edwards, 1837 ) ( = Uca lactea perplexa) — Fukui, Wada and Wang, 

1989; Tzeng and Chen, 1992; Shih, 1994; Wang and Liu, 1996a, b; Yu, Jeng, Chan, Ho and 

Shy, 1996; Jeng, 1997, 1998; Wang and Liu, 1998; Lee and Tung, 2000; Shih, 2000a 

Uca tetragonon (Herbst, 1790) — Ho, Wang, Lin and Yu, 1993; Shih, 1994; Wang and Liu, 

1996a, b; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997,1998; Wang and Liu, 1998; Lee 

and Tung, 2000; Shih, 2000a 

Uca triangularis (A. Milne Edwards, 1873) — Wang, 1984; Dai, Yang, Song and Chen, 1986;


Fukui, Wada and Wang, 1989; Huang, Yu and Takeda, 1989; Dai and Yang, 1991; Huang, 

1994; Shih, 1994; Wang and Liu, 1996b; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997, 

1998; Lee and Tung, 2000; Shih, 2000a 

Subfamily Macrophthalminae Dana, 1851 * r t,i 

Macrophthalmus abbreviates Manning and Holthuis, 1981 ( = Macrophthalmus dilatatus (De Haan, 

1835)) — Maki and Tsuchiya, 1923; Takahasi, 1934a, b, 1935; Sato, 1936b, c, d; Sakai, 

1939, 1940; Chang, 1963; Sakai, 1976; Dai, Yang, Song and Chen, 1986; Fukui, Wada and 

Wang, 1989; Dai and Yang, 1991; Shih, Lue and Wang, 1991; Huang, Yu and Takeda, 1992; 

Kuo, 1995; Hsueh, 1996; Wang and Liu, 1996a, b; Ho and Hung, 1997; Jeng, Lin, Wang and 

Jan, 1998; Lin, 1998; Wang and Liu, 1998 

Macrophthalmus banzai Wada and Sakai, 1989 ( = Macrophthalmus japonicus aut. ) — Maki and 

Tsuchiya, 1923; Takahasi, 1935; Sakai, 1939; Lin, 1949; Wu, Ling, Shieh and Wang, 1962; 

,W0 Chang, 1963; Sakai, 1976; Su and Lue, 1984; Dai, Yang, Song and Chen, 1986; Wada and 

Sakai, 1989; Dai and Yang, 1991; Shih, Lue and Wang, 1991; Ho, 1995; Kuo, 1995; Hsueh, 

1996; Wang and Liu, 1996a, b; Ho and Hung, 1997; Jeng, Lin, Wang and Jan. 1998; Lin, 

1998; Wang and Liu, 1998; Jeng and Wang, 2000; Lee and Tung, 2000 

Macrophthalmus boscii Audouin, 1826 — Sakai, 1939, 1940; Horikawa, 1940; Lin, 1949; Sakai, 

1976; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, Yu and Takeda, 1992; 

Huang, 1994; Jeng, 1997 

Macrophthalmus boteltobagoe Sakai, 1939 — Sakai, 1939; Horikawa, 1940; Sakai, 1940; Lin, 

1949; Sakai, 1976; Huang, Yu and Takeda, 1992; Huang, 1994 

Macrophthalmus ceratophorus Sakai, 1969 — Ho, 1995 [Fig. 8b] 

Macrophthalmus convexus Stimpson, 1858 — Maki and Tsuchiya, 1923; Sakai, 1939; Sakai, 1976; 

WJ3 Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, Yu and Takeda, 1992; Huang, 

1994 

Macrophthalmus latreillei (Desmarest, 1822)—present record [Fig. 8c] 

Macrophthalmus quadratus A. Milne Edwards, 1873 — Jeng, 1997 

Macrophthalmus serenei Takeda and Komai, 1991 ( = Macrophthalmus verreauxi aut ) — Jeng, 1997 

Macrophthalmus tomentosus Souleyet, 1841 — Balss, 1922c; Sakai, 1940 

Subfamily Scopimerinae Alcock, 1900 

llyoplax delsmani De Man, 1926 — Takahasi, 1935 [Fig. 8d] [This species has not been collected 

since Takahasi's (1935) report, and this is rather surprising considering the amount of work on 

ocypodids which has been conducted in Taiwan over the last decade. Of the various llyoplax 

species, /. delsmani in particular, is very distinctive, with courting males ivory white in colour 

(see Ng and Sivasothi, 1999) and it is very unlikely that it could have missed by workers. We retain 

Takahasi's record for the moment, but with some doubt. ] 

llyoplax formosensis Rathbun, 1921 — Rathbun, 1921; Maki and Tsuchiya, 1923; Takahasi, 1932, 

1934a, b; Sakai, 1939, 1940; Wu, Ling, Shieh and Wang, 1962; Sakai, 1976; Dai, Yang, 

Song and Chen, 1986; Fukui, Wada and Wang, 1989; Dai and Yang, 1991; Huang, Yu and 

Takeda, 1992; Huang, 1994; Wang and Liu, 1996a, b; Wang and Liu, 1998 [This species was


Taiwan, but Kitaura, Wada and Nishida (1998) recently reported it from 

northern Vietnam. Takahasi's (1932) record of this species from Taiwan may contain specimens of 

/. tansuiensis as well (Sakai, 1939). ] 

Ityoplax Integra (Tesch, 1918) —Sakai, 1939, 1940, 1976; Huang, Yu and Takeda, 1992; Wang 

and Liu, 1996a, b; Yu, Jeng, Chan, Ho and Shy. 1996; Jeng, 1997; Kitaura, Wada and 

Nishida, 1998; Wang and Liu, 1998 

Ityoplax tansuiensis Sakai, 1939 — Sakai, 1939, 1940, 1976; Dai, Yang, Song and Chen, 1986; 

Fukui, Wada and Wang, 1989; DaiandYang, 1991; Huang, Yuand Takeda, 1992; Huang, 1994; 

Hsueh, 1996; Wang and Liu, 1996a, b; Ho and Hung, 1997; Jeng, Lin, Wang and Jan, 1998; 

Kitaura, Wada and Nishida, 1998; Wang and Liu, 1998 

Scopimera bitympana Shen, 1930 —Takahasi, 1935; Sato, 1936d; Sakai, 1939, 1940, 1976; Dai, 

Yang, Song and Chen, 1986; Fukui, Wada and Wang, 1989; DaiandYang, 1991; Huang, Yu 

and Takeda, 1992; Huang, 1994; Hsueh, 1996; Wang and Liu, 1996a, b; Ho and Hung, 1997; 

Jeng, 1997; Jeng, Lin, Wang and Jan, 1998; Lin, 1998; Kitaura, Wada and Nishida, 1998; 

Wang and Liu, 1998; Jeng and Wang, 2000; Lee and Tung, 2000 

Scopimera globosa De Haan, 1835 — Takahasi, 1934a, b, 1935; Sato, 1936a, b; Sakai, 1939; 

Chang, 1963; Sakai, 1976; Dai, Yang, Song and Chen, 1986; Fukui, Wada and Wang, 1989; ^ f l ^ 

DaiandYang, 1991; Shih, Lue and Wang, 1991; Huang, Yu and Takeda, 1992; Huang, 1994; ' 

Kuo, 1995; Ho and Hung, 1997; Lee and Tung, 2000 

Scopimera longidactyla Shen, 1932 — Sakai, 1940, 1976; Dai, Yang, Song and Chen, 1986; 

Fukui, Wada and Wang, 1989; DaiandYang, 1991; Huang, Yu and Takeda, 1992; Hsueh, 

1996; Wang and Liu, 1996b; Ho and Hung, 1997; Lee and Tung, 2000 

Tmethypocoelis ceratophora (Koelbel, 1897)— Takahasi, 1935; Sakai, 1940, 1976; Dai, Yang, 

Song and Chen, 1986; Fukui, Wada and Wang, 1989; DaiandYang, 1991; Shih, Lue and 

Wang, 1991; Huang, Yu and Takeda, 1992; Wu, 1992; Huang, 1994; Kuo, 1995; Wang and 

Liu. 1996a, b; Ho and Hung, 1997; Jeng, 1997; Jeng, Lin, Wang and Jan, 1998; Wang and 

Liu, 1998; Jeng and Wang, 2000; Lee and Tung, 2000 

Family Camptandriidae Stimpson, 1858 

Baruna sinensis Tan and Huang, 1995 ( = Leipocten sordidulum aut. )—-Sakai, 1939, 1940, 1976; 

Dai, Yang, Song and Chen, 1986; Fukui, Wada and Wang, 1989; DaiandYang, 1991; Huang, 

Yu and Takeda, 1992; Huang, 1994; Tan and Huang, 1995: Hsueh, 1996 

Paracleistostoma depressum De Man, 1895 — Fukui, Wada and Wang, 1989; Huang, Yu and Takeda, 

1992; Kitaura, Wada and Nishida, 1998 

Family Mictyridae Dana, 1852 

Mictyris brevidactylus Stimpson, 1858 ( = Mictyris tongicarpus aut. ) — Parisi, 1918; Balss, 1922c; .^-^i 

Maki and Tsuchiya, 1923; Takahasi, 1934a, 1934b, 1935; Sato, 1936a, b, c, d; Sakai, 1939; 

Horikawa, 1940; Lin, 1949; Chang, 1963; Sakai, 1976; Su and Lue, 1984; Dai, Yang, Song 

and Chen, 1986; Fukui, Wada and Wang, 1989; DaiandYang, 1991; Shih, Lue and Wang, 

1991; Huang, Yu and Takeda, 1992; Wu, 1992; Huang, 1994; Kuo, 1995; Hsueh, 1996; 

Wang and Liu, 1996a, b; Ho and Hung, 1997; Jeng, Lin, Wang and Jan, 1998; Lin, 1998;



Family Grapsidae MacLeay, 1838 

Subfamily Grapsinae MacLeay, 1838 

Geograpsus crinipes (Dana, 1851)—Wang and Liu, 1996a; Ho, 1997; Wang and Liu, 1998; Liu, 

1999 

Geograpsus grayi (H. Milne Edwards, 1853) —Sakai, 1939; Horikawa, 1940; Lin, 1949; Sakai, 

1976; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994; Wang and Liu, 

1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Ho, 1997; Jeng, 1997; Wang and Liu, 1998; Liu, 

1999 

Geograpsus stormi De Man, 1895— Jeng, Jan, Tzeng, Feng and Yang, 1996; Ho, 1997; Jeng, 

Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Liu, 1999 

Grapsus albolineatus Lamarck, 1818 ( = Grapsus strigosus aut. )—Oshima. 1921a; Balss, 1922c; 

Maki and Tsuchiya, 1923; Sakai, 1939; Hwang and Yu, 1980, Wang, 1984; Suzuki, 1985; 

Dai, Yang, Song and Chen, 1986; Fukui, Wada and Wang, 1989; Dai and Yang, 1991; Chang 

and Chen, 1992; Tzeng and Chen, 1992; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 

1994, 1996; Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Ho and Hung, 1997; 

Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Lai, Huang and Fang, 1997; Jeng, Shao, 

Tzeng, Feng and Wu, 1998; Ng, 1998; Wang and Liu. 1998; Lee and Tung, 2000 

Grapsus longitarsis Dana, 1851 — Sakai, 1939; Horikawa, 1940; Lin, 1949 

Grapsus tenuicrustatus (Herbst, 1783) — Maki and Tsuchiya, 1923; Sakai, 1939; Hwang and Yu, 

1980; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Tzeng and Chen, 1992; Chang 

and Chen, 1992; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 1994; Ho, 1996; Jeng, Jan, 

Tzeng, Feng and Yang, 1996; Wang and Liu, 1996a; Jeng, Jan, Tzeng and Feng, 1997; Lai, 

Huang and Fang, 1997; Jeng, Lin, Wang and Jan, 1998; Jeng, Shao, Tzeng, Feng and Wu, 

1998; Ng, 1998; Wang and Liu, 1998; Lee and Tung, 2000 

Metopograpsus messor (Forskal, 1775 ) — Balss, 1922c; Hiro, 1939; Sakai, 1939; Wu, Ling, 

Shieh and Wang, 1962; Chang, 1963; Sakai, 1976; Hwang and Yu, 1980; Wang, 1984; 

Fukui, Wada and Wang, 1989; Chang and Chen, 1992; Jeng, Jan, Tzeng, Feng and Yang, 

1996; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998 

Metopograpsus thukuhar (Owen, 1839) — Fukui, Wada and Wang, 1989; Chang and Chen, 1992; 

Hsueh, 1996; Jeng, Jan, Tzeng, Feng and Yang, 1996; Wang and Liu, 1996a, b; Yu, Jeng, 

Chan, Ho and Shy, 1996; Ho and Hung, 1997; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; 

Lai, Huang and Fang, 1997; Huang and Liitzen, 1998; Jeng, Lin, Wang and Jan, 1998; Jeng, 

Shao, Tzeng, Feng and Wu, 1998; Wang and Liu, 1998; Jeng and Wang, 2000; Lee and Tung, 

2000 

Packygrapsus crassipes Randall, 1839 — Sato, 1936c, d; Tzeng and Chen, 1992 

Pachygrapsus fakaravensis Rathbun, 1907 — Hwang and Yu, 1980 [Fig. 8e] [The species is apparently 

not common in East Asia, being more common in the central Pacific. The specimen figured is 

from Hawaii, where it was also only recently reported. Although superficially similar to P. plicatus 

in general morphology, their live colours are very different, with P. fakaravensis much darker 

overall. ]


Paehygrapsus minutus A. Milne Edwards, 1873 — Sakai, 1939; Suzuki, 1985; Dai, Yang, Song 

and Chen, 1986; Dai and Yang, 1991; Chang and Chen, 1992; Huang, 1994; Jeng, Jan, 

Tzeng, Feng and Yang, 1996; Wang and Liu, 1996a; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 

1997; Lai, Huang and Fang, 1997; Huang and Liitzen, 1998; Jeng, Shao, Tzeng, Feng and 

Wu, 1998; Wang and Liu, 1998 

Paehygrapsus plicatus (H. Milne Edwards, 1873) —Huang, 1994; Wang and Liu, 1996a; Lai, 

Huang and Fang, 1997; Wang and Liu, 1998 

Planes cyaneus Dana, 1851 ( = Planes minutus aut. ) — Sakai, 1939; Dai, Yang, Song and Chen, 

1986; Fukui, Wada and Wang, 1989; Dai and Yang, 1991 ^ fc|


1991; Wu, 1992; Huang, 1994; Hsueh, 1996; Jeng, Jan, Tzeng, Feng and Yang, 1996; Wang 

and Liu, 1996b; Ho and Hung, 1997; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, 

Shao, Tzeng, Feng and Wu, 1998 [Ng and Liu (1998) has suggested that the genus Chiromantes 

be restricted to the type species, C. haematocheir, and the other species, including C. dehaani, 

should be referred to other genera. Studies by P. K, L, Ng and C. D. Schubart are now ongoing 

to revise the genus Chiromantes. ] 

Clistocoeloma sinense Shen, 1933 ( = Cleistocoeloma merguiense aut. ) — Sakai, 1939; Dai, Yang, 

Song and Chen, 1986; Fukui, Wada and Wang, 1989; Dai and Yang, 1991; Hsueh, 1995; 

Hsueh, 1996; Hsueh and Huang, 1996a 

Episesarma lafondU (Jacquinot, in Hombron and Jacquinot, 1843) ( = Episesarma mederi aut. ) — 

Ng, 1998 [Ng (1998), who examined a female specimen collected by H. -C. Liu from southern 

Taiwan identified it as Episesarma mederi (H. Milne Edwards, 1854). The first author recently 

examined specimens of E. lafondU from Japan and the Taiwanese specimen should be referred to 

E. lafondU instead] 

Labuanium rotundatum (Hess, 1865)—Sakai, 1939, 1940, 1976; Dai, Yang, Song and Chen, 

^ 1986; Dai and Yang, 1991; Huang, 1994; Liu, 1999 [This interesting species was originally re- 

^ ^ ported from Tanshui in northern Taiwan by Sakai (1939 ) but we have been unable to find the 

species there, probably because the area has been extensively developed over the last 50 years. In 

wiifW 

stead, we have since found this very agile species in southern Taiwan, in Kenting, living in 

tree-holes filled with water (phytotelms). They spend almost all their time on trees, and have 

been observed several metres from the ground. Females come to the ground more frequently on 

their way to spawn in the sea which may be some distance away. Adult specimens are coffee-brown 

to dirty yellow with younger specimens more mottled. ] 

Metasesarma aubryi A. Milne Edwards, 1869 — Maki and Tsuchiya, 1923; Sakai, 1939, 1940, 

1976; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Huang, 1994; Liu, 1999 

Metasesarma rousseauxi H. Milne Edwards, 1853 ( = Measesarma granulans Heller, 1862 ) — 

Balss, 1922c; Liu, 1999 [This supralittoral species (see Ng and Davie, 1995) has surprisingly 

not been reported from Taiwan since Balss, but there are a good number of specimens in the 

TMCD and ZRC from the islands of Lanyu and Lutao as well as Kenting National Park] 

Nanosesarma gordoni (Shen, 1935)—Fukui, Wada and Wang, 1989; Yu, Jeng, Chan, Ho and 

Shy, 1996; Jeng, 1997 [This species may well be a junior synonym of N. minutum (De Man, 

1887) as has been suggested (P. J. F. Davie, personal communication) but until both taxa are 

formally synonymised, we prefer to keep them species separate for the time being. ] 

Nanosesarma minutum (De Man, 1887) — Hsueh, 1996; Lai, Huang and Fang, 1997 

Neosarmatium fourmanoiri Serene, 1973 — Ng, Liu and Wang, 1997 

Neosarmatium indicum (A. Milne Edwards, 1868)-Ng, Liu and Wang, 1997 

Neosarmatium meinerti (De Man, 1887)—Horikawa, 1940; Sakai, 1941; Lin, 1949; Dai, Yang, 

Song and Chen, 1986; Dai and Yang, 1991; Davie, 1994; Ng, Liu and Wang, 1997 

Neosarmatiumpunctatum (A. Milne Edwards, 1873) — Ng, Liu and Wang, 1997 

Neosarmatium rotundifrons (A. Milne Edwards, 1869) — Ng, Liu and Wang, 1997 

Parasesarma acis Davie, 1993 ( = Sesarma erythrodactylum aut. ) — Suzuki, 1985; Fukui, Wada 

r-^ and Wang, 1989 [This species is almost certainly synonymous with Parasesarma tripectinis 

K'"5f«c

Ng et al. : Checklist of urabs from Taiwan 43 

(Shen, 1940) (Dai, A. -Y. personal communication) and will be dealt with shortly by her in 

a paper on the Hong Kong fauna. Until this is formally done, we retain the use of this name. ] 

Parasesarma pictum (De Haan, 1835 ) ( = Sesarma pictum) — Balss, 1922c; Maki and ,*. , 

Tsuchiya, 1923; Sato, 1936c, d; Sakai, 1939; Chang, 1963; Sakai, 1976; Su and Lue, 4 '^ 

1984; Dai, Yang, Song and Chen, 1986; Fukui, Wada and Wang, 1989; Dai and Yang, 

1991; Shih, Lue and Wang, 1991; Chang and Chen, 1992; Tzeng and Chen, 1992; Wu, 

1992; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 1994; Hsueh, 1996; Jeng, Jan, 

Tzeng, Feng and Yang, 1996; Wang and Liu, 1996b; Ho and Hung, 1997; Jeng, Jan, 

Tzeng and Feng, 1997; Jeng, Lin, Wang and Jan, 1998; Jeng, Shao, Tzeng, Feng and 

Wu, 1998; Wang and Liu, 1998; Jeng and Wang, 2000; Lee and Tung, 2000 

Parasesarma plicatum (Latreille, 1806) (= Ocypode (Pachysoma) affinis De Haan, 1837; Sesarma 

plicata) — Maki and Tsuchiya, 1923; Sakai, 1939; Su and Lue, 1984; Wang, 1984; 

Dai, Yang, Song and Chen, 1986; Fukui, Wada and Wang, 1989; Dai and Yang, 1991; 

Shih, Lue and Wang, 1991; Wu, 1992; Huang, 1994; Hsueh, 1995; Kuo, 1995; Hsueh, 

1996; Jeng, Jan, Tzeng, Feng and Yang, 1996; Wang and Liu, 1996b; Ho and Hung, 

1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Lin, Wang and Jan, 1998; Jeng, Shao, 

Tzeng, Feng and Wu, 1998; Lin, 1998; Wang and Liu, 1998; Lee and Tung, 2000 

[Ongoing studies by P. K. L. Ng and A. -Y. Dai indicate that Ocypode (Pachysoma) ajfinis De 

Haan, 1837 (type locality Japan) is a separate species from Parasesarma plicatum (Latreille, 

1806) (type locality East India) and should be regarded as distinct. These studies indicate that 

the Taiwanese specimens which have been referred to " Parasesarma plicatum" clearly belong 

to P. affinis instead. For the time being, and until these results are published, we use the 

name Parasesarma plicatum solely for convenience] 

Parasesarma tripectinis (Shen, 1940) — Hsueh, 1995, 1996 [See comments for Parasesarma 

acis above] 

Perisesarma bidens (De Haan, 1835 ) ( = Sesarma bidens) — Oshima, 1921a; Maki and -ftfL-L 

Tsuchiya, 1923; Sato, 1936d; Sakai, 1939; Wu. Ling, Shieh and Wang, 1962; Sakai, 

1976; Dai, Yang, Song and Chen, 1986; Fukui, Wada and Wang, 1989; Dai and Yang, 

1991; Shih, Lue and Wang, 1991; Huang, 1994; Hsueh, 1995, 1996; Jeng, Jan, Tzeng, Feng 

and Yang, 1996; Wang and Liu, 1996a, b; Yu, Jeng, Chan, Ho and Shy, 1996; Ho and 

Hung, 1997; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Lin, Wang and Jan, 

1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Wang and Liu, 1998; Lee and Tung, 2000 

Sesarmops impressum (H. Milne Edwards, 1837) (= Sesarma impressum) — Sakai, 1939, 

1940, 1976; Wang, 1984; Dai, Yang, Song and Chen, 1986; Fukui, Wada and Wang, 

1989; Dai and Yang, 1991; Huang, 1994; Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and 

Shy, 1996; Jeng, 1997, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Wang and Liu, 1998; 

Liu, 1999 

Sesarmops intermedium (De Han, 1835)—Terao, 1916; Balss, 1922c; Sakai, 1939; Sakai, 4nVy«fcf, 

1976; Hwang and Yu, 1980; Wang, 1984; Fukui, Wada and Wang, 1989; Wang and Liu, 

1996a; Yu, Jeng, Chan, HoandShy, 1996; Jeng, 1997, 1998; Wang and Liu, 1998; Liu, 1999 

Stelgistra stormi De Man, 1895— Ng and Liu, 1999

44 National Taiwan Museum Special Publication Series, No. 11, 86 pp,, April 2001 

Subfamily VaruninaeH. MilneEdwards, 1853 

Acmaeopleura parvula Stimpson, 1858 — Fukui, Wada and Wang, 1989; Jeng, Shao, Tzeng, Feng and 

Wu, 1998 

Chasmagnathus convexus De Haan, 1835 — Terao, 1916; Maki and Tsuchiya, 1923; Sato, 1936a, d; 

Sakai, 1939; Wu, Ling, Shieh and Wang, 1962; Sakai, 1976; Su and Lue, 1984; Dai, Yang, Song 

and Chen, 1986; Fukui, Wada and Wang, 1989; Dai and Yang, 1991; Shih, Lue and Wang, 1991; 

Wu, 1992; Huang, 1994; Hsueh, 1996; Wang and Liu, 1996a, b; Ho and Hung, 1997; Jeng, Lin, 

Wang and Jan, 1998; Lin, 1998; Wang and Liu, 1998; Lee and Tung, 2000 

Cyclograpsus integer H. MilneEdwards, 1837 —Fukui, Wada and Wang, 1989 

Cyclograpsus intermedius Ortmann, 1894 — Sakai, 1976; Dai, Yang, Song and Chen, 1986; Dai 

and Yang, 1991; Huang, 1994 [In the TMCD are specimens of this species - 1 male, 1 female 

(TMCD2362), Hsiaoliuchiu Island, Pingtung County, coll. J.-T. Lin, 17 May 1988; 1 female 

(TMCD 2478), Keelung, coll. L.-C. Sun, 2 January 1989; 3 males, 4 females (TMCD 2557), 

Hsiaohsianglan, Fulung, Taipei County, coll. C.-H. Wang, 21 April 1989; 1 female (TMCD 

2565), Keelung, coll. L.-C. Sun, 7 May 1989. ] 

Eriocheir japonica De Haan, 1835 ( = Eriocheir rectus Stimpson, 1858, following Chan, Hung and 

Yu, 1995) —Terao, 1915a; Oshima, 1921a; Balss, 1922c; Maki and Tsuchiya, 1923; Sato, 

1936c, d; Sakai, 1939; Wu, Ling, Shieh and Wang, 1962; Chang, 1963; Chang, 1965; Sakai, 

1976; Wang and Chen, 1981; Su and Lue, 1984; Dai, Yang, Song and Chen, 1986; Lai, Shy 

F»/W3 and Yu, 1986; Yu and Ho, 1986; Fukui, Wada and Wang, 1989; Dai and Yang, 1991; Shih, 

Lue and Wang, 1991; Chang and Chen, 1992; Hung, Ho and Yu, 1992; Wu, 1992; Jeng, Jan, 

Tzeng, Feng and Yang, 1994; Ho, 1996; Hsueh, 1996; Jeng, Jan, Tzeng, Feng and Yang, 

1996; Wang and Liu, 1996b; Guo, Ng, Dai and Ng, 1997; Ho and Hung, 1997; Jeng, Jan, 

Tzeng and Feng, 1997; Jeng, Lin, Wang and Jan, 1998; Jeng, Shao, Tzeng, Feng and Wu, 

1998; Shy and Yu, 1999; Chan, 2000 [Nakagawa (1915a, b) used a new name Eriocheir 

formosa for a species from Taiwan but this name is a nomen nudum (see discusssion for Platyeriocheir 

formosa). Nakagawa ( 1915a, b), however, did note that this was only a provsional 

nomenclature. Koba (1936) indicates that Nakagawa's name is actually a synonym of Eriocheir 

japonica. ] 

Gaetice depressus (De Haan, 1833) — Maki and Tsuchiya, 1923; Sato, 1936c, d; Sakai, 1939, 

1940; Lin, 1949; Dai, Yang, Song and Chen, 1986; Fukui, Wada and Wang, 1989; Dai and 

IfyA/flt? Yang, 1991; Shih, Lue and Wang, 1991; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 

1994; Hsueh, 1996; Jeng, Jan, Tzeng, Feng and Yang, 1996; Wang and Liu, 1996a, b; Yu, 

Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Lin, 

Wang and Jan, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Wang and Liu, 1998; Jeng and 

Wang, 2000; Lee and Tung, 2000 

Helice formosensis Rathbun, 1931 ( = Helice tridens aut. part, Helice tridens latimera aut. part) — 

oW,l1*? Rathbun, 1931; Parisi, 1918; Maki and Tsuchiya, 1923; Sakai, 1939; Horikawa, 1940; Saki, 

1940; Chang, 1963; Sakai and Yatsuzuka, 1980; Dai, Yang, Song and Chen, 1986; Fukui, 

Wada and Wang, 1989; Dai and Yang, 1991; Shih, Lue and Wang, 1991; Huang, 1994; 

Hsueh, 1995; Kuo, 1995; Hsueh, 1996; Wang and Liu, 1996a, b, c; Ho and Hung, 1997; 

Jeng, Lin, Wang and Jan, 1998; Lin, 1998; Wang and Liu, 1998; Jeng and Wang, 2000; Lee

Ng el al. : Checklist of crabs from Taiwan 

and Tung, 2000 [Fig. 8f] [The records of H. tridens (not Ocypode ( Helice) tridens De Haan, 

1835) by Terao (1916), Sato (1936c, d), Horikawa (1940), Wu, Ling, Shieh and Wang 

(1962), Chiu (1964), Sakai (1940, 1976), Su and Lue (1984), Shih, Lue and Wang (1991) 

and Wu (1992) cannot be accurately ascertained, many probably being mixed species records including 

H. formosensis and/or Helice sp. (see later). Similarly, records of H. latimera (not 

Helice tridens latimera Parisi, 1918) from Taiwan by Maki and Tsuchiya (1923), Sakai (1939, 

1940, 1976), Wu, Ling, Shieh and Wang (1962) and Wang and Liu (1996c, from Kinmen) are 

probably also H. formosensis (M. Tiirkay, personal communication to P. K. L. Ng). See also 

comments for Helice sp. later on ] 

Helice subquadrata (Dana, 1851) (= Helice leachii Hess, 1865) — Sakai, 1939; Sakai, 1976; 

Dai, Yang, Song and Chen, 1986; Fukui, Wada and Wang, 1989; Dai and Yang, 1991; Shih, 

Lue and Wang, 1991; Huang, 1994; Hsueh, 1996; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 

1997; Lee and Tung, 2000 [Fig. 8g. ] [The old records of H. leachii from many parts of the 

Indo-West Pacific are actually H. subquadrata according to M. Tiirkay (personal communication 

to P. K. L. Ng), and this species will also be referred to a new genus when the revision of this 

group is completed, see comments for Helice sp. later on] 

Helice sp. (= Helice tridens aut. part; Helice wuana nut. part) — Sakai, 1939, 1940, 1976; Sakai 

and Yatsuzuka, 1980; Dai, Yang, Song and Chen, 1986; Fukui, Wada and Wang, 1989; Dai 

and Yang, 1991; Shih, Lue and Wang, 1991; Huang, 1994; Hsueh, 1996; Wang and Liu, 

1996a, b; Ho and Hung, 1997; Jeng, Lin, Wang and Jan, 1998; Lin, 19988; Wang and Liu, 

1998; Jeng and Wang, 2000; Lee and Tung, 2000 [Fig. 8h] [The specimens reported by various 

authors as Helice wuana (not Helice tridens wuana Rathbun, 1931) from Taiwan will be referred 

to a new species in a soon to be finished revision of the genera Chasmagnathus and Helice by K. 

Sakai, M. Tiirkay and S.-L. Yang (M. Tiirkay, personal communication to P. K. L. Ng). He 

also comments that the records of H. sheni Sakai, 1939, by Sakai (1939, 1976) (including 

records from Taiwan) are also H. wuana s. str., the two names being subjective synonyms. Dai 

and Yang's (1991) and Hsueh's (1995, 1996) records of H. japonica (not Helice {Helicana) 

japonica K. Sakai and Yatsuzuka, 1980) are also likely to belong to the present new species. Helice 

wuana (and the present new species) was referred to the subgenus Helicana Sakai and Yatsuzuka, 

1980, by these authors and in the upcoming revision, Helicana will be regarded as a distinct 

genus. ] 

Hemigrapsus penicillatus (De Haan, 1835 ) — Sakai, 1939, 1940; Wu, Ling, Shieh and Wang, . ^ , ..„„ 

1962; Sakai, 1976; Dai, Yang, Song and Chen, 1986; Fukui, Wada and Wang, 1989; Dai and ^ ' 

Yang, 1991; Shih, Lue and Wang. 1991; Hsueh, 1996; Wang and Liu, 1996a, b; Ho and 

Hung, 1997; Jeng, Lin, Wang and Jan, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Wang 

and Liu, 1998; Chou, Lai and Fang, 1999; Jeng and Wang, 2000; Lee and Tung, 2000 

Hemigrapsus sanguineus (DeHaan, 1835) — Balss, 1922c; Sakai, 1939; Sakai, 1976; Dai, Yang, 

Song and Chen, 1986; Fukui, Wada and Wang, 1989; Dai and Yang, 1991; Hsueh, 1996; + ****' 

Jeng, Jan, Tzeng, Feng and Yang, 1996; Wang and Liu, 1996a, b; Ho and Hung, 1997; Jeng, 

Jan, Tzeng and Feng, 1997; Jeng, Lin, Wang and Jan, 1998; Jeng, Shao, Tzeng, Feng and 

Wu, 1998; Wang and Liu, 1998; Jeng and Wang, 2000 

Metaphtx elegans De Man, 1888 — Fukui, Wada and Wang, 1989; Shih, Lue and Wang, 1991; 

45


Kuo, 1995; Wang and Liu, 1996a, b; Ho and Hung, 1997; Jeng, Lin, Wang and Jan, 1998; 


Metaplax takahashii Sakai, 1939 — Sakai, 1939, 1940, 1976; Dai, Yang, Song and Chen, 1986; 

Fukui, Wada and Wang, 1989; Dai and Yang, 1991 

Parapyxtdognathus deianira (De Man, 1888)—Fukui, Wada and Wang, 1989 

Platyeriocheirformosa (Chan, Hung and Yu, 1995) (= Eriocheir rectus aut., Eriocheirformosa) — 

Sakai, 1938b; Sakai, 1939, 1940; Lin, 1949; Sakai, 1976; Wang and Chen, 1981; Dai, 

Yang, Song and Chen, 1986; Yu and Ho, 1986; Dai and Yang, 1991; Hung, Ho and Yu, 1992; 

Shy and Yu, 1992; Huang, 1994; Chan, Hung and Yu, 1995; Ho, 1996; Hung and Yu, 1996; 

Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997, 1998; Ng, Guo and Ng, 1999; Shy and Yu, 

1999; Chan, 2000 [The name Eriocheir formosa, as used by Chan, Hung and Yu (1995), has 

actually been used much earlier by Nakagawa (1915a, b), but this name is a nomen nudum as 

there was no accompanying description, figure or any indication. As such, this name is not 

nomenclaturally available. Koba (1936) indicates that Eriocheir formosa Nakagawa, 1915a, is 

actually conspecific with Eriocheir japonica. ] 

Pseudograpsus atbus Stimpson, 1858 — Suzuki, 1985 

Pseudograpsus setosus (Fabricius, 1778) — Ng, Jeng and Ng, in press 

Ptychognathus barbatus (A. Milne Edwards, 1873) — Maki and Tsuchiya, 1923; Fukui, Wada and 

Wang, 1989; Chang and Chen, 1992; Huang, 1994; Jeng, Jan, Tzeung, Feng and Yang, 1996; 

Wang and Liu, 1996a, b; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng 

and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Wang and Liu, 1998 

Ptychognathus hachijyoensis Sakai, 1955 — Fukui, Wada and Wang, 1989 

Ptychognathus ishii Sakai, 1939 —Sakai, 1939, 1940, 1976 

Ptychognathus takahasii Sakai, 1939 — Sakai, 1939, 1940, 1976 

Thalassograpsus harpax (Hilgendorf, 1892) —Sakai, 1939, 1940, 1976; Dai, Yang, Song and 

Chen, 1986; Dai and Yang, 1991 

Utica borneensis De Man, 1895 — Maki and Tsuchiya, 1923; Horikawa, 1940; Lin, 1949; Dai, 


Varuna litterata (Fabricius, 1798) —Balss, 1922c; Maki and Tsuchiya, 1923; Sakai, 1939; Wu, 

Ling, Shieh and Wang, 1962; Wang and Chen, 1981; Dai, Yang, Song and Chen, 1986; 

Hwang and Takeda, 1986; Fukui, Wada and Wang, 1989; Dai and Yang, 1991; Tzeng and 

Chen, 1992; Huang, 1994; Ho, 1996; Hsueh, 1996; Wang and Liu, 1996a, b; Yu, Jeng, 

Chan, Ho and Shy, 1996; Ho and Hung, 1997; Jeng, 1997; Lai, Huang and Fang, 1997; Jeng, 

1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng, 1998; Wang and Liu, 1998; Shy and Yu, 

1999; Lee and Tung, 2000 [Many of the older records of this species should be rechecked as some 

might have been mistaken identifications for the very closely related V. yui (see Ng, 1998). 

Varuna litterata, however, does seem to be the more common species in Taiwan. ] 

Varuna yui Hwang and Takeda, 1986 — Hwang and Takeda, 1986; Huang, 1994; Jeng, Jan, Tzeng, 

Feng and Yang, 1996; Jeng, Jan, Tzeng and Feng, 1997; Jeng, Shao, Tzeng, Feng and Wu, 

1998; Ng, 1998 

Xenograpsus testudinatus Ng, Huang and Ho, 2000 — Ng, Huang and Ho, 2000 [Through the courtesy 

of M. -S. Jeng, the first author and N. K. Ng examined a very large series of this species,


including very small individuals. The differences between this taxon and the closely allied X. 

novaeinsularis (from a different arc of volcanic islands) noted by the original authors remain valid, 

except perhaps for the convexity of the carapare which may be difficult to discern in small specimens. 

Ongoing studies also indicate that Xenograpsus occupies a very isolated position in the 

Varuninae (and Grapsidae) and its systematic position must be reappraised. ] 

Family Plagusiidae Dana, 1851 

Percnon abbreviatum (Dana, 1851)—Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997 

Percnon planissimum (Herbst, 1804) ( = Percnon tenuifrons aut. ) — Balss, 1922c; Maki and 

Tsuchiya, 1923; Sato, 1936d; Sakai, 1939; Hwang and Yu, 1980; Wang, 1984; Chang and 

Chen, 1992; Tzeng and Chen, 1992; Jeng. Jan, Tzeng. Feng and Yang, 1994, 1996; Wang and 

Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1997; Jeng, Jan, Tzeng and Feng, 1997; 

Lai, Huang and Fang, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Wang and Liu, 1998; 

Lee and Tung, 2000 

Plagusia dentipes De Haan, 1335 ( = Plagusia chabrus aut. ) — Sakai, 1939, 1976; Dai, Yang, 

Song and Chen, 1986; Dawson, 1987; Dai and Yang, 1991; Huang, 1994 [Maki and Tsuchiya 

(1923) reported Plagusia chabrus (Linnaeus, 1758) from Taiwan but his figure clearly shows /*. 

dentipes instead. Plagusia chabrus is not yet known from the northern hemisphere (Dawson, 

1987 ) . In the National Taiwan Museum are two dried specimens (male, 57. 6 by 49. 5 mm 

(TMCD1572), female, 51. 0 by 46. 8 mm (TMCD 1571), Hopingtao, Keelung, coll. S.-H. 

Wu, August 1996). Unlike other Plagusia species in Taiwan, P. dentipes is sublittoral, living at 

depths up to 30 m deep and as such, not often seem (S. -H. Wu, personal communication). ] 

Plagusia immaculata Lamarck, 1818 — present record [The presence of this species in Taiwan was 

only recently confirmed (see Schubart and Ng, 2000). It bears a very close resemblance to P. 

squamosa but adults can most easily be separated by the degree of setatation on the carapace and 

chelipeds. ] 20o^ 

Plagusia speciosa Dana, 1852 — Ng, Liu and Ho. \n presfr [This interesting species was only recently 

reported from southern Taiwan. 3 

Plagusia squamosa (Herbst, 1790) ( = Plagusia tuberculata Lamarck, 1818; Plagusia depressa 

aut. ) — Oshima, 1921a; Balss, 1922c; Maki and Tsuchiya, 1923; Sato, 1936d; Takahasi, 

1936c; Sakai, 1939; Hwang and Yu, 1980; Dai, Yang, Song and Chen, 1986; Dawson, 1987; 

Dai and Yang, 1991; Huang, 1994; Jeng, Jan, Tzeng, Feng and Yang, 1994; Ho, 1996; Jeng, 

Jan, Tzeng, Feng and Yang, 1996; Wang and Liu, 1996a; Ho and Hung, 1997; Lai, Huang and 

Fang, 1997; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng, 1998; Wang and Liu, 1998; Lee and 

Tung, 2000 [This species is best known in the literature as P. tuberculata or P. depressa tuberculata 

but a recent study by Schubart, Gonzalez-Gordillo, Reyns, Liu and Cuesta (2001) provides 

evidence that it should be regarded as a distinct species; and P. squamosa (Hersbt, 1790) is the 

most senior name (Schubart and Ng, 2000). Another name, Plagusia orientalis Stimpson, 1858 

(type locality Hong Kong), is also a junior snynonym (Schubart and Ng, 2000). ] 

Family Pinnotneridae De Haan, 1833 

Subfamily Pinnotherinae De Haan, 1833 

47


Pinnotheres boninensis Stimpson, 1858 —JLHO and Hung, 1997; Lee and Tung, 2000 

Pinnotheresparvulus Stimpson, 1858 — Maki, 1931: Horikawa, 1940 

Pinnotheres taichungae Sakai, 2000 ( = Pinnotheres bidentatus aut. ) — Hsueh and Huang, 1996b; 

Sakai, 2000 [Sakai (2000) felt taht Pinnotheres bidentatus reported by Hsueh and Hung (19%) 

from Taiwan was a misidentification and recently created a new name for them, Pinnotheres 

i taichungae. 

f ; k Pinnotheres tsingtaoensis Shen, 1932 — Soong, 1997 

Subfamily Asthenognathinae Stimpson, 1858 

Asthenognathus sp. — Lin, 1949 [The identity of this species is a problem as Lin's specimen(s) are 

no longer extant. According to Sakai (1976: 588), the only species in East Asian waters is the 

supposed Japanese endemic Asthenognathus inaequipes Stimpson, 1858. Serene and Soh (1976) 

added a third species from me Indo-Wesl Pacific and recorded A. hexagonum Rathbun, 1909, 

from the Philippines. Fn view of the proximity of Taiwan to Japan and the Philippines, Lin's specimen(s) 

might well belong to either A. inaequipes or A. hexagonum. ] 

Subfamily Xenophthalminae Alcock, 1900 

Neoxenophthalmus obscurus (Henderson, 1893) — Fang, 1991; Huang, 1994 

Family Cryptochiridae Paulson, 1875 

Hapalocarcinus marsupialis Stimpson, 1858 — Utinomi, 1944; Yu, Jeng, Chan, Ho and Shy, 

1996; Jeng, 1997, 1998 

Pseudocryptochirus viridis Hiro, 1938 — Utinomi, 1944 

Family Palicidae Rathbun, 1898 

Subfamily Palicinae Rathbun. 1898 

Pseudopalicus oahuensis (Rathbun, 1906) —Castro, 2000 

Subfamily Crossotonotinae Moosa and Serene, 1981 

Crossotonotus spinipes (DeMan, 1888) —Ng, Chan and Wang, 2000; Castro, 2000 

Family Gecarcinidae MacLeay, 1838 

Cardisoma carnifex (Herbst, 1794) —Oshima, 1921a; Balss, 1922c; Maki and Tsuchiya, 1923; 

Sakai, 1939; Lin, 1949; Tiirkay, 1974; Sakai, 1976; Wang and Chen, 1981; Dai, Yang, Song 

and Chen, 1986; Dai and Yang, 1991; Ho, Shy and Yu, 1992; Huang, 1994; Kuo, 1995; 

Wang and Liu, 1996a, b; Yu, Jeng, Chan, Ho and Shy, 1996; Ho and Hung, 1997; Jeng, 

1997, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Lin, 1998; Ng, 1998; Wang and Liu, 

1998; Liu, 1999; Lee and Tung, 2000 

Discoplax hirtipes (Dana, 1851) (= Cardisoma hirtipes) — Sakai, 1939; Tiirkay, 1974; Sakai, 

P> 1976; Dai, Yang, Song and Chen, 1986; Dai and Yang, 1991; Ho, Shy and Yu, 1992; Huang, 

J 1994; Wang and Liu, 1996a; Yu, Jeng, Chan, Ho and Shy, 1996; Ho, 1997d; Ho and Hung,


1997; Jeng, 1997, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Ng, 1998; Wang and Liu, 

1998; Liu, 1999 [The generic placement of this species and D. rotundum follows Tiirkay (1987) 

who noted that the genus Discoplax A. Milne Edwards, 1867 (not 1873 as often cited), was 

valid, see also Ng, Nakasone and Kosuge (2000). A reappraisal of this genus and its differences 

with Cardisoma sensu stricto is now being undertaken by D. Guinot and P. K. L. Ng, with molecular 

studies of the family by C. D. Schubart and his associates. ] 

Discoplax rotundum (Quoy and Gaimard, 1824 ) ( = Cardisoma rotundum) — Ho, Shy and Yu, 

1992; Ng, 1998; Liu, 1999 

Epigrapsus notatus (Heller, 1865) —Ng, Liu and Wang, 1999; Liu, 1999 

Gecarcoidea lalandii H. Milne Edwards, 1837 — Sakai, 1939, 1940, 1976; Dai, Yang, Song and ^fflt)^ i$? 

Chen, 1986; Dai and Yang, 1991; Ho, Shy and Yu, 1992; Huang, 1994; Wang and Liu, 

1996a; Yu, leng, Chan, Ho and Shy, 1996; Jeng, 1997, 1998; Jeng, Shao, Tzeng, Feng and 

Wu, 1998; Ng, 1998; Wang and Liu, 1998; Liu, 1999 

Family Potamidae Ortmann, 1896 

The Sinopotamidae Bott, 1970, and Isolapotamidae Bott, 1970, are now regarded as junior synonyms 

of the Potamidae (Ng and Dudgeon, 1992; Ng and Trontelj, 1996; Dai, 1997; Ng and Tan, 

1998). 

Candidiopotamon rathbunae (De Man, 1914 ) ( = Candidiopotamon rathbuni) — De Man, 1914; 

Oshima, 1921a; Balss, 1922c; Maki and Tsuchiya, 1923; Balss, 1937; Sakai, 1939, 1940; 

Pretzmann, 1963; Bott, 1967; Bott, 1970; Sakai, 1976; Wang, 1984; Hwang and Mizue, 

1985; Ng, 1996b; Shy, Lai and Yu, 1996; Yu, Jeng, Chan, Ho and Shy, 1996; Jeng, 1998; 

Jeng, Lin, Wang and Jan, 1998; Dai, 1999; Ng, 1999b; Shy and Yu, 1999; Lee and Tung, 

2000; Shih, 2000b [Most authors spell the specific epithet "rathbuni", which is incorrect. Although 

De Man (1914) named this species " Potamon (Potamon) rathbuni", he also wrote that 

"Cette jolie espece que j'ai l'honneur de dedierau savant auteur de la belle Monographic des Crabes 

d'eau douce, parue il y a quelques annees dans les "Nouvelles Archives du Museum ..." (De 

Man, 1914: 128). The author of this important book, of course, was Mary Rathbun, a lady. In 

the manner De Man originally spelled the name, however, it implied that the species is named after 

a man, which is clearly not the case here. Article 32c (ii) of the current International Code of Zoological 

Nomenclature (1999), states that if "... in the original publication itself, without recourse 

to any external source of information, clear evidence of an inadvertent error ... ", then the name 

can be emended. As this is clearly the situation in this case, the species name of De Man's taxon 

should be corrected to Candidiopotamon rathbunae instead (see also Ng, 2000a). There is also an 

additional nomenclatural matter with regards to a long forgotten name. Nakagawa (1915a, b) used 

a new scientific name " Thelphusa rubra" for a Taiwanese crab for which he also applied a Taiwanese 

vernacular name. There was no description, figure or indication and such, the name Thelphusa 

rubra Nakagawa, 1915a, is clearly a nomen nudum. In any case, from the Taiwanese vernacular 

name used, we have little doubt that Thelphusa rubra Nakagawa, 1915a, is actually Candidiopotamon 

rathbunae. This species is also often coloured dark to almost bright red, easily explaining 

the specific name (" rubra'') chosen by Nakagawa (1915a, b). ]


Geothelphusa albogilva Shy, Ng and Yu, 1994 ( = Nanhaipotamon formosanum aut. ) — Suzuki, 

1985; Shy, Ng and Yu, 1994; Shy, Lai and Yu, 1996; Yu, Jeng, Chan, Ho and Shy, 1996; 

Jeng, 1997, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Dai, 1999; Liu, 1999; Shy and 

Yu, 1999; Shih, 2000b 

Geothelphusa ancylophallus Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Shy, Lai and Yu, 

1996; Dai, 1999; Shy and Yu, 1999; Lee and Tung, 2000 

Geothelphusa bicolor Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 

1999 

Geothelphusa caesia Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Liu, 1999; Shy 

andYu, 1999 

Geothelphusa candidiensis Bott, 1967 (= Geothelphusa dehaani mi. ) — Bott, 1967; Bott, 1970; 

Minei, 1974; Hwang and Mizue, 1985; Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 1999; 

Shih, 2000b [Oshima (1921a) reported " Potamon {Geothelphusa) dehaanii candidiensis Rath- 

bun" from Taiwan. This name, however, has never been formally published by Rathbun as far as 

we known. This name as appears in Oshima (1921a) must thus be regarded as nomen nudum, as 

there was no acccompanying description. Records supposedly of this species from the Ryukyus in 

Japan (see Sakai, 1939, 1976) almost certainly belong to other species (Shy, Ng and Yu, 1994), 

with the record from Ishigaki recently referred to a new species (Shy and Ng, 1998). ] 

Geothelphusa chiui Minei, 1974 — Minei, 1974; Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 

1999 [Records by Hwang and Mizue (1985 ) of this species must be regarded as doubtful and 

should be rechecked (see Shy, Ng and Yu, 1994). See also discussion for G. olea. ] 

Geothelphusa cinerea Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 

1999 

Geothelphusa dolichopodes Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and 

Yu, 1999 

Geothelphusa eucrinodonta Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and 

Yu, 1999 

Geothelphusa eurysoma Shy, Ng and Yu, 1994 —Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 

1999 

Geothelphusa ferruginea Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Yu, Jeng, Chan, Ho and 

Shy, 1996; Jeng, 1997, 1998; Jeng, Shao, Tzeng, Feng and Wu, 1998; Dai, 1999; Shy and 

Yu, 1999 

Geothelphusa gracilipes Shy, Ng and Yu, 1994 —Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 

1999 

Geothelphusa hirsuta Tan and Liu, 1998 — Tan and Liu, 1998; Shy and Yu, 1999 

Geothelphusa ilan Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 1999 

Geothelphusa lanyu Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 

1999 

Geothelphusa lutao Shy, Ng and Yu, 1994 —Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 1999 

Geothelphusa miyazakii (Miyake and Chiu, 1965) ( = Geothelphusa dehaani aut. ) — Miyake and 

Chiu, 1965; Minei, 1974; Hwang and Mizue, 1985; Shy, Ng and Yu, 1994; Dai, 1999; Shy


and Yu, 1999 [Records of this species from the Ryukyus in Japan (see Sakai, 1976 ) almost 

certainly belong to other species (Shy, Ng and Yu, 1994). ] 

Geothelphusa monticola Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 

1999 

Geothelphusa nanao Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 

1999 

Geothelphusa nanhsi Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Shy, Lai and Yu, 1996; 

Dai, 1999; Shy and Yu, 1999; Lee and Tung, 2000 

Geothelphusa neipu Chen, Cheng and Shy, 1998 — Chen, Cheng and Shy, 1998; Ng, 1999b; Shy 

and Yu, 1999; Shih. 2000b; Shy, Chen and Cheng, 2000 [This species is very close to G. ping- 

tung and may be synonymous. The colour differences observed between these two species (Shy, 

Chen and Cheng, 2000) are not always reliable as both colour form, supposedly diagnostic for 

each of these two species, have recently been collected together. In any case, the type localities of 

both species are very close to each other] 

Geothelphusa olea Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Shy, Lai and Yu, 1996; Dai, 

1999; Liu, 1999; Shy and Yu, 1999; Lee and Tung, 2000 [ Geothelphusa olea has a very wide 

distribution in western Taiwan, and its presently known range actually includes the type locality of 

G. chiui Minei, 1974, a species whose identity has been the subject of some discussion (see Shy, 

Ng and Yu, 1994). Geothelphusa chiui has not been found from its type locality inNanpu (Hsinchu 

County) despite numerous efforts (H. -C. Liu and J. -Y. Shy, personal communication) and de 

spite a recent search, the male holotype cannot be found in the Kitakyushu Museum in Japan where 

the rest of Minei's material now resides (D. C J. Yeo, personal communication). Shy, Ng and 

Yu (1994) chose to recognise G. olea and G. chiui as separate taxa because they discerned some 

differences in the form of the male first pleopods between the specimens of G. olea they had on 

hand and of G. chiui using the figures in Minei's original 1974 paper. The extent of recent collec 

tions of Taiwanese freshwater crabs, which covers a very wide area, suggests that these differences 

may not be valid. The absence of any Geothelphusa species resembling G. chiui or G. olea from 

the type locality of G. chiui may just be because the area has been too extensively developed and 

the population there has become extinct. It seems rather unlikely that two closely related species 

(i.e. G. olea and G. chiui) are living in close proximity or even sympatric with each other; more 

likely both are synonyms. However, we retain the use of both names until this can be confirmed. ] 

Geothelphusa pingtung Tan and Liu, 1998—Tan and Liu, 1998; Shy and Yu, 1999 

Geothelphusa takuan Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 

1999 

Geothelphusa tali Shy, Ng and Yu, 1994 —Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 1999 

Geothelphusa taroko Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 

1999 

Geothelphusa tawu Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 1999 

Geothelphusa tsayae Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 

1999; Shih, 2000b 

Geothelphusa wangi Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 

1999


Geothelphusa wutai Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and Yu, 

1999 

Geothelphusa yangmingshan Shy, Ng and Yu, 1994 — Shy, Ng and Yu, 1994; Dai, 1999; Shy and 

Yu, 1999 

Nanhaipotamon formosanum (Parisi 1916) — Parisi. 1916; Balss, 1922c, 1937; Sakai, 1939, 

1940; Bott, 1968a, 1970; Sakai, 1976; Hwang and Mizue, 1985; Dai, 1997; Dai, 1999; Shy 

andYu, 1999; Lee and Tung, 2000 

NB. Records of Geothelphusa dehaani by Terao ( 1915a), Balss ( 1922c), Maki and Tsuchiya 

(1923), Balss (1937), Sakai (1939, 1940), Lin (1949), Wu, Ling, Shieh and Wang (1962) 

and Wang and Chen (1981), cannot be determined; most likely all their records are mixed and belong 

to one of the above Geothelphusa species instead. The real G. dehaani is endemic to the 

Japanese main island. 

Family Parathelphusidae Alcock, 1910 

Somanniathelphusa taiwanensis Bott, 1968 ( = Parathelphusa sinensis aut. ) — Terao, 1915b; 

Balss, 1922c; Maki and Tsuchiya, 1923; Sakai, 1939, 1940; Bott, 1968b; Bott, 1970; Hwang 

and Mizue, 1985; Shy, Lai and Yu, 1996; Dai, 1999; Shy and Yu, 1999; Lee and Tung, 2000 

Endemic taxa 

From the conservation perspective, there are a number of species which are endemic to Taiwan. 

Foremost in consideration (and concern) must surely be the true freshwater crabs, notably Candidiopotamon 

rathbunae, Geothelphusa albogilva, G. ancylophallus, G. bicolor, G. caesia, G. candidiensis, 

G. chiui, G cinerea, G. dolichopodes, G eucrinodonta, G. eurysoma, G. ferruginea, 

G. gracilipes, G. hirsuta, G. ilan, G lanyu, G lutao, G. miyazakii, G. monticola, G. nanao, G. 

nanhsi, G neipu, G. olea, G pingtung, G. takuan, G tali, G. taroko, G tawu, G tsayae, G. wangi 

G. wutai, G. yangmingshan, Nanhaipotamon foimosanum (Potamidae), and Somanniathelphusa 

taiwanensis (Parathelphusidae) (Ho, 1998a-e). All 34 species are endemic to Taiwan. Records of G. 

candidiensis and G. miyazakii from the Ryukyus actually belong to other species (see Shy and Ng, 

1998). Of these, the widespread Candidiopotamon rathbunae is not seriously threatened by extinction, 

but most of the other species have very narrow distributions and easily be exterminated by large scale developments(Shih, 

2000b). 

The Taiwanese or green hairy crab ( Platyeriocheir formosa, Grapsidae) is known only from eastern 

Taiwan, and Platyeriocheir is the only known endemic Taiwanese brachyuran genus. Although 

generally regarded as a freshwater species, the species has larvae which develop in the open sea, and is 

a very vulnerable species as its adult habitat requirements are very specific (Hung and Yu, 1996; Chan, 

2000 ). Requiring clean montane streams, the species is extremely vulnerable to pollution and/or 

over-exploitation by collectors - female crabs are much sought after in the breeding season because of 

their ovaries. 

The large marine Taiwanese fiddler crab ( Uca formosensis, Ocypodidae) interestingly, has been 

found only in Taiwan so far (including Penghu), and has not been reported from nearby China, Philippines 

or Ryukyus, and may well be Taiwan's only marine endemic crab. Its conservation is now a mat-

]Ng et al. : Checklist of crabs from Taiwan 53 

ter of some concern (Shih, 1997; Shih, Mok, Chang and Lee, 1999; Shin, 2000a). The overgrowth 

of mangroves and development projects are serious threats to the long term survival of this species. 

Two other wholly marine species, Dumeia taiwanicus and Acanthonyx formosa (Majidae), Tymolus 

hirtipes Tan and Huang, 2000 (Cyclodorippidae), Helice formosensis Rathbun, 1931 , 

Xenograpsus testudinatus Ng, Huang and Ho, 2000 (Grapsidae), and Pinnotheres taichungae Sakai, 

2000 (Pinnotheridae, see discussion for this species) now known only from Taiwan, but are probably 

more widespread than they are now once more intensive surveys are carried out. One other species, 

Jonas choprai (Corystidae), is formally known only from Taiwan, but it has been found in the Indian 

Ocean (P. K. L. Ng, unpublished data). 

In addition to the above-mentioned taxa, the following species fravc- •IGO^ described from Taiwan, 

but have been found in other parts of the world, viz. Leucosia formosensis Sakai, 1937 (Leucosiidae), 

Calappa quadrimaculata Takeda and Shikatani, 1990 (Calappidae), Jonas formosae (Balss, 1922) 

(Corystidae), Demania rotundata Serene, in Guinot, 1969 (Xanthidae), Pilumnus acanthosoma Ng, 

2000 (Pilumnidae), Trapezia cheni Galil, 1983 , Trapezia garthi Galil, 1983 (Trapeziidae), 

Chaceon manningi Ng, Lee and Yu, 1994 (Geryonidae), Macrophthalmus boteltobagoe Sakai, 

1939, Hyoplax formosensis Rathbun, 1921, Hyoplax tansuiensis Sakai, 1939 (Ocypodidae), Baruna 

sinensis Tan and Huang, 1995 (Camptandriidae), and Ptychognathus ishii Sakai, 1939, Ptychognathus 

takahasii Sakai, 1939, Varuna yui Hwang and Takeda, 1986 (Grapsidae). One described 

species, Eurcrate formosensis Sakai, 1974, is now regarded as a junior synonym of E. alcocki Serene, 

in Serene and Lohavanijaya, 1973, although it may be resurrected as a valid taxon in the future (see 

discussion for this species). One species, Elamena truncata (Stimpson, 1858) (Hymenosomatidae), 

originally described from the Ryukyus in southern Japan, had its type locality changed to northern Taiwan 

when Ng and Chuang (1996) designated a neotype from there to stabilise the taxonomy of this and 

allied species. 

On the fauna from Kinmen 

The brachyurans of Kinmen have been most recently reported on by Yang and Chang (1996) and 

Wang and Liu (1996c). Between them, they record a total of 47 species, viz. Dromiidae: Lauridromia 

dehaani; Dorippidae: Paradorippe granulata; Leucosiidae: Philyra pisum; Calappidae: Calappa 

lophos, C. phitargius; Matutidae: Matuta planipes, M. victor (as M. lunaris): Orithyiidae: 

Orithyia sinica; Majidae: Doclea japonica, Hyasienus diacanthus; Parthenopidae: Platylambrus 

validus, Cryptopodia fornicata; Corystidae: Jonas distincta; Portunidae: Portunus gracilimanus, 

Portunus haanii, P. pelagicus, P. sanguinolentus, P. trituberculatus, Scyllaparamamosain (as S. serrata), 

Charybdis feriatus, C. japonica, C. natator, Thalamita crenata; Xanthidae: Leptodius nigromaculatus, 

Atergatis reticulatus; Pilumnidae: Heteropanope glabra; Goneplacidae: Carcinoplax 

longimana; Ocypodidae: Ocypode ceratophthalmus, O. sinensis (as O. cordimana), Ilea arcuata, U. 

borealis, V. lactea, Scopimera tuberculata, S. longidactyla, Macrophthalmus abbreviatus (as M. dilatatus), 

M. banzai (as M. japonicus), M. convexus, M. erato; Mictyridae: Mictyris brevidactylus; 

and Grapsidae: Metopograpsus quadridentatus, Hemigrapsus penicillatus, Varuna litterata, Helice sp. 

(as Helice wuana), Metaplax longipes, Parasesarmapictum, P. plicatum and Perisesarma bidens. 

Of these species, only the following are unique to this area and not yet reported from the main island 

of Taiwan itself, viz. Orithyia sinica (Linnaeus, 1771) (Orithyiidae), Leptodius nigromaculatus


Serene, 1962 (Xanthidae), Heteropanope glabra Stimpson, 1858 (Pilumnidae), Macrophthalmus erato 

De Man, 1888, Scopimera tube reulata Stimpson, 1858 (Ocypodidae), Metopograpsus quadridentatus 

De Man, 1858, and Metaplax longipes Stimpson, 1858 (Grapsidae). Of these, notable is also 

that the family Orithyhdae is only known from Kinmen as far as Taiwanese territory is concerned. 

On the fauna from Tungsha Islands 

Ng, Lee and Yu (1994) described Chaceon manningi (Geryonidae) from the Tungsha Islands. Ng 

and Huang (1997) subsequently recorded Paromola japonica Parisi, 1915, P. macrochira Sakai, 

1961 (Homolidae) and Platymaia bartschi Rathbun, 1916, and Cyrtomaia curviceros Bouvier, 1915 

(Majidae) from these islands. Most recently, Ho and Ng (1999 ), Ng and Chen (1999) and Ng 

(2000b) recorded Homolodromia kai Guinot, 1993 (Homolodromiidae), Lamoha longirostris (Chen, 

1986) (Homolidae) and Benthochascon hemingi Alcock and Anderson, 1899 [Fig. 4f] (Portunidae), 

respectively from Tungsha. We here add two more species from deep waters off these islands, viz. 

Paromolopsis boasi Wood-Mason and Alcock, 1891 (Homolidae) [Fig. lbjand Platymaia fimbriata 

Rathbun, 1916 (Majidae) [Fig. 4b] . Of these species Homolodromia kai, Lamoha longirostris, 

Paromolopsis boasi, Cyrtomaia curviceros, Platymaia fimbriata and Benthochascon hemingi are 

known only from Tungsha thus far and have not been recorded in Taiwan itself. Their presence in Taiwan 

proper, however, must be expected as all of them have wide distributions which bracket the island. 

General discussion 

A total of 548 species from 36 families are now known from the main island of Taiwan as defined 

here. This figure compares very favourably with the number now known from neighbouring China and 

Japan (cf. Dai and Yang, 1991; Sakai, 1976), especially considering the small size of the island. 

Some families, notably the Leucosiidae, Majidae, Xanthidae, Pilumnidae, Goneplacidae, Pinnotheridae 

and Cryptochiridae, are still poorly represented for the Taiwanese fauna. Not surprisingly, 

many are small taxa with cryptic habits (e.g. Cryptochiridae) and/or their taxonomy is very difficult 

(e. g. Leucosiidae, Pilumnidae). For example, only six and two species of Pinnotheridae and Cryptochiridae 

respectively are known from the island, a ridiculously small number. For a very speciose 

group like the Xanthidae, many more new records can be expected once the deeper waters and the coral 

reefs are more intensely explored. Even for the well studied swimming crabs (Portunidae), we know of 

a good number of unpublished new records, including species of Carupa, Charybdis and Ovalipes 

(P. -H. Ho and H. -C. Liu, personal communication). We are also aware of substantial numbers of 

new records of Goneplacidae (J. -F. Huang and P. -W. Hsueh, personal communication) and Majidae 

(S. -H. Wu and T. -Y. Chan, personal communication) which are now being studied or written up. 

Even in the intertidal area, where a great deal of work has been done, there are still many surprises. In 

the Grapsidae, there is at least one new genus and several new species now being studied by the first author, 

C. D. Schubart and H. -C. Liu. In addition, there are new records of Grapsus, Chiromantes, 

(Grapsidae) and Epigrapsus (Gecarcinidae) (C. D. Schubart and H. -C. Liu, personal communication). 

Several families are still conspicuously absent from the Taiwanese brachyuran fauna thus far, viz. 

Homolodromiidae, Cymonomidae. Phyllotymolinidae, Orithyiidae, Retroplumidae and Hexapodidae. 

We believe that these will probably be discovered in Taiwan once the proper in depth surveys are


conducted. Hexapodids are small crabs which can be easily missed in collections and are often mistaken 

for damaged goneplacids. In fact, Huang (1994) reports Hexapus anfractus (Rathbun, 1910)froman 

unspecified site in the Taiwan Straits without stating from which side if the strait it was from, but its 

presence in Taiwan itself must be expected. Cymonomids, phyllotymolinids and retroplumids are 

deep-water crabs and are difficult to collect, but should be present in the deep waters off eastern and 

southern Taiwan. 

The Orithyiidae Dana, 1852, is a peculiar group normally found in shallow waters along continental 

China and Korea, and for Taiwan, it is only known from Kinmen, near China (Yang and Chang, 

1996) thus far. Its absence from the main island of Taiwan itself is striking, although it is possible it 

may just be very rare there. The Homolodromiidae Alcock, 1899, only known from deep waters off the 

Tungsha Islands thus far (Ho and Ng, 1999), is also certainly present near the main island of Taiwan as 

well. 

From what we now know being studied by ourselves and various associates, the crab fauna for Taiwan 

will easily reach 600 species. We believe that once most of the surveys and studies have been completed, 

the total brachyuran fauna for Taiwan and its associated islands will probably approach 700. 

These figures compare very well with the total brachyuran fauna of the world which now stands at over 

6000 species. Considering that Taiwan's relatively small size and the fact that it has some 10% of the 

world's crab species, this is a very impressive fauna indeed. All in all, the future for carcinological discoveries 

in Taiwan is excellent, and there is still some way to go before its fauna can be considered to be 

well known! 

Acknowledgements 

The present study was supported by the National Taiwan Museum (Taipei), and provided two travel 

fellowships to the first author. The first author was also partially supported by a research grant from 

the National University of Singapore. Part of the third author's work was supported by a grant to Professor 

Hsiang-Ping Yu (National Taiwan Ocean University) from the National Science Council of Taiwan. 

We are most appreciative of the generous help rendered to us by Dr. Hung-Chang Liu (National Tsing 

Hua University), Mr. Jun-Tsong Lin (National Taiwan Museum), Dr. Jung-Fu Huang (National 

Kaohsiung Institute of Marine Technology), Dr. Jhy-Yun Shy (c/o National Taiwan Ocean University), 

Dr. Pan-Wen Hsueh (National Chung Hsing University), Mr. Ding-An Lee (The Taiwan Fisheries 

Research Institute) ans Mr. Kuan-Hsing Li (Peikuan Crab Museum) who have helped make this 

study possible. Dr. Hui-Lian Chen (Oceanological Institute, Chinese Academy of Science, Qingdao) 

was very kind in helping identify some of the leucosiids. We are also very grateful to Professor 

Hsiang-Ping Yu (National Taiwan Ocean University) for his support and kind help throughout this project; 

as well as to Professor Tin-Yam Chan (National Taiwan Ocean University) for his assistance in 

many aspects of the present study. The first author is very grateful to Mr. Paul Clark for his advice on 

the correct authors and dates for the species described by Jacquinot, Adams and White; Dr. Michael 

Tiirkay (Senckenberg Museum), Mr. Peter Davie (Queensland Museum), Prof. Ai-Yun Dai 

(Academia Sinica, Beijing), Dr. Hui-Lian Chen (Oceanological Institute, Qingdao), Dr. Christoph 

Schubart, Ms. Ng Ngan Kee, Mr. Tan Swee Hee and Mr, Darren Yeo (National University of Singapore) 

for sharing their expertise on the groups they are now studying with him, and their many useful 

suggestions, which have helped keep this list as current as possible.


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Appendix 1 

New records from Taiwan and Tungsha Islands 

1. Homola mieensis Sakai, 1979 [Fig. la] 

Material examined — 1 female, 36. 0 by 42. 6 mm (TMCD), Tahsi, Ilan County, 400 m, coll. S. -H. 

Wu, 20 April 1995; 1 female, 37. 2 by 41. 4 mm (NTOU 9003-02-29), Tungsha Islands, 600 m, 

coll. D. -A. Lee, 22 February 1990 

2. Paromolopsis boasi Wood-Mason and Alcock, 1891 [Fig. lb] 

Material examined — 1 female, 32. 5 by 33. 1 mm (NTOU), Tungsha Islands, coll. P. -H. Ho, 21 

April 1995 

3. Notosceles serratifrons (Henderson, 1893) [Fig. lc] 

Material examined — 1 dried female, 15. 4 by 29. 8 mm (TMCD 394), Fangliao, Pingtung County, 

coll. C. -Y. Wei, 7 December 1969 

4. Leucosia rhomboidalis De Haan, 1850 [Fig. If] 

Material examined — 1 male, 15. 4 by 17. 8 mm (TMCD 259), Tainan, coll. C. -Y. Wei, 20 October 

1963; 1 dried male, 16. 1 by 19. 0 mm (TMCD 232), Kaohsiung, coll. C.-Y. Wei, 1 November 

1968; 1 male (TMCD 2812), Yungan, Kaohsiung County, coll. K.-C. Hsia 

5. Arcania septemspinosa (Fabricius, 1787) [Figs. 2a, b] 

Material examined — 1 male, 38. 4 by 22. 8 mm (TMCD 408), Anping fish market, Tainan City, 

coll. C. -Y. Wei, 10 December 1964 

6. Ixa edwardsii Lucas, 1858 [Figs. 2d, e] 

Material examined — 1 dried female, 69. 6 by 33. 4 mm (TMCD 363), Taiwan, coll. C.-Y. Wei, 14 

October 1971; 1 dried female, 74. 8 by 39.1 mm (TMCD 284), Pingtung, coll. C.-Y. Wei, 25 January 

1968 

7. Myra biconica Ihle, 1918 [Fig. 2f] 

Material examined — 1 dried female, 24. 3 by 32.1 mm (TMCD 419), Tainan, coll. C.-Y. Wei, 18 

December 1967 

8. Nursiaplicata (Herbst, 1803) [Fig. 2g] 

Material examined — 1 dried female, 21. 7 by 16. 5 mm (TMCD 406), Anping Harbour, Tainan City, 

coll. C. -Y. Wei, 10 December 1975; 1 dried female, 20. 9 by 16. 3 mm (TMCD 407), Anping 

Harbour, Tainan City, coll. C. -Y. Wei, 10 December 1975 

9. Pariphiculus mariannae (Herklots, 1852) [Fig. 2h] 

Material examined — 1 dried male, 25. 4 by 29. 7 mm (TMCD 220), Fangliao, Pingtung County, 

coll. C. -Y. Wei, 16 February 1962; 1 dried male, 27. 5 by 32. 2 mm (TMCD 436 ), Hengchun,

Pingtung County, coll. C. -Y. Wei, 20 August 1970 


10. Philyra heterograna Ortmann, 1892 [Fig. 3a] 

Materia] examined — 1 dried male, 17. 6 by 18. 1 mm (TMCD 427), Kaohsiung, coll. C.-Y. Wei, 2 

July 1983 

11. Catappa bicornis Miers, 1884 [Fig. 3b] 

Material examined — 1 dried female, 32. 0 by 24. 6 mm (TMCD 419), Tainan, coll. C.-Y. Wei, 18 

December 1967; 1 female, 99. 7 by 75. 8 mm (NTOU), Hopingtao, Keelung City, coll. S.-H. Wu, 

May 1998; 1 male (ZRC), Makang, Han County, coll. 1996 

12. Cyrtomaia curviceros Bouvier, 1915 [Fig. 3h] 

Material examined — 1 dried female, 45. 1 by 47. 5 mm (spines inclusive) (Peikuan Crab Museum), 

Tahsi, Ilan County, coll. Peikuan Museum, 2000. 

13. Platymaia bartschi Rathbun, 1916 [Fig. 4a] 

Material examined—1 male specimen (TMCD 2226), Tahsi, Ilan County, coll. J.-T. Lin, 22 January 

1987 

14. Platymaia fimbriata Rathbun, 1916 [Fig. 4b] 

Material examined — 1 female (ZRC), Tungsha Islands, coll. P.-H. Ho, 14 January 1995 

15. Pleistacantha sanctijohannis Miers, 1879 [Fig. 4c] 

Material examined — 1 dried male, 14. 8 by 19. 3 mm (TMCD 454), Taitung, coll. C. -Y. Wei, 15 

October 1964; 1 dried female (15. 6 by 22. 9 mm) (TMCD 455), Taitung, coll. C.-Y. Wei, 15 October 

1964 

16. Lissocarcinus arkati Kemp, 1923 [Fig. 4e] 

Material examined — 1 dried male, 35. 4 by 21. 6 mm (TMCD 479), Keelung fish market, Keelung 

City, coll. C.-Y. Wei, 12 March 1974; 1 dried female, 20. 9 by 17. 5 mm (TMCD 480), Keelung 

fish market, Keelung City, coll. C.-Y. Wei, 12 March 1974; 1 dried female, 25.3 by 21.1 mm 

(TMCD 200), Kaohsiung, coll. C.-Y. Wei, 20 January 1971 

17. Carcinoplax indica Doflein, 1904 [Fig. 7c] 

Material examined — 1 male, 30. 1 by 23. 0 mm (ZRC 1998. 216), Tahsi, Ilan County, coll. P. K. L. 

Ng, August 1996; 1 male, 30. 9 by 24. 8 mm, 1 female. 35. 0 by 27. 3 mm (ZRC 1999. 773), Tahsi, 

Ilan County, coll. P. K. L. Ng and K. Lim, May 1999 

18. Carcinoplax tomentosa Sakai, 1969 [Fig. 7e] 

Material examined — 1 male, 45. 7 by 34. 9 mm (TMCD 1587), Tahsi, Ilan County, coll. S. -H. 

Wu, 27 December 1995; 1 male, 45. 6 by 34. 3 mm (TMCD 1653), Tahsi, Ilan County, coll. S.-H. 

Wu, March 1996; 1 female (TMCD 2228), Tahsi, Ilan County, coll. J. -T. Lin, 22 January 1987; 1


male, 53. 5 by 40. 7 mm, 1 female, 46. 3 by 35. 3 mm (ZRC 1998.43Mater4 ), Nangfangao, Ilan 

County, coll. S. H. Tan, 28 May 1998; 1 male, 29. 3 by 22. 9 mm (ZRC 1999. 720), Tungkang, 

Pingtung County, coll. P. K. L. Ng and K. Lim, May 1999; 1 female (ZRC 1997. 748), coll. P. 

K. L. Ng, 3 September 1996 

19. Eucrate Solaris Yang and Sun, 1979 [Fig. 7h] 

Material examined — 1 female, 26. 2 by 21. 1 mm (ZRC 1997. 752), Tungkang, Pingtung County, 

coll. P. K. L. Ng, 5 August 1996 

20. Macrophthalmus latreillei (Desmarest, 1822) [Fig. 8c] 

Materia] examined—1 male, 38. 8 by 28. 6 mm, (TMCD), Szutsao, Tainan City, coll. J.-H. Lee, May 

1998; 1 female, 45. 7 by 33. 1 mm (TMCD), Szutsao, Tainan City, coll. J.-H. Lee, April 16, 2000

Romanization 

Anping 

Changhua County 

Diaoyutai 

Fangliao 

Fulung 

Hengchun 

Homei 

Hopingtao 

Hsiangshan 

Hsiaohsianglan 

Hsiaoliuchiu 

Hsinchu City 

Ilan County 

Kaohsiung City 

Kaohsiung County 

Keelung City 

Kenting National Park 

Lanyu 

Lukang 

Lutao 

Makang 

Nangfangao 

Nanliao 

Penghu 

Pingtung County 

Pratas Islands 

Suao 

Szutsao 

Tahsi 

Tainan City 

Taipei County 

Taitung County 

Tanshui 

Tiaoyutai 

Toucheng 

Tungkang 

Tungsha Islands 

Yungan 


Appendix 2 

Taiwanese localities (Chinese) and their romanizations as used in the text 

Chinese 

iCT (in Tainan City) 

$!&£(= Tiaoyutai) 

^jjfF (in Pingtung County) 

11^ (in Taipei County) 

H^ (in Pingtung County) 

fTJ3! (in Taipei County) 

frffJlGn Keelung City) 

If |i[ (in Hsinchu City) 

/Mffli (in Taipei County) 

/hlW (in Pingtung County) 

mi 

m 

STBIt^&llil (in Pingtung County) 

ftftfg (in Taitung County) 

fH*?§Kin Changhua County) 

$kM (in Taitung County) 

Hift (in Ilan County) 

j^f^ft (in Ilan County) 

j?J^ (in Kaohsiung County) 

mm 

M&Mik ( = Tungsha Islands, in Kaohsiung City ) 

MM (in Ilan County) 

K^ (in Tainan City) 

XM (in Ilan County) 

^^K (in Taipei County) 

$}1&M.( = Diaoyutai) 

Sl^ (in Ilan County) 

^i# (in Pingtung County) 

%&%$% ( = Pratas Islands, in Kaohsiung City) 

JK^Z (in Kaohsiung County)


Appendix 3 

Journal names in Chinese or Japanese and their English translations as used in the text 

English 

Annuals of Taiwan Museum 

Aquiculture 

Biological Bulletin of National Taiwan Normal University 

Biological Bulletin of the Tunghai University 

Botany and Zoology 

CHYUGAIIZISHINPOU 

Fisheries Extension, Taipei 

Journal of Fisheries Society of Taiwan 

Journal of Oceanography in Taiwan Strait 

Oceanologia et Limnologia Sinica 

Quarterly Journal of Natural Conservation 

Scientific Taiwan 

Taiwan Natural Science 

Transactions of the Natural History Society of Formosa 

Zoological Magazine 

ZYUZENKAIZASHI 

Chinese 

wm$m 

mmsm


< > 

Fig. 1. a. Homola mieensis, female (NTOU). Tungsha; b. Paromolopsis boasi, female (NTOU), 

Tungsha; c Notosceles serratifrons, dried female (TMCD394), Fangliao [preserved coloration]: 

d. Dorippe sinica, female (ZRC 1999. 470). Guangdong, southern China [freshly 

preserved coloration]; e. Leucosia haematosticta, male (ZRC), Singapore; f, Leucosia 

rhomhoidalis, dried male (TMCD259). Tainan [preserved coloration]; g, Leucosia vittata, 

male (ZRC). Thailand; h, Leucosia anatum. mate (ZRC). Singapore. 

79


Fig. 2. a, Arcania septemspinosa, dried male (TMCD 408), Tainan [preserved coloration]; b, Arcania 

septemspinosa, male (ZRC), Thailand; c, Ixacylindrus, female (ZRC), Thailand; d, 

Ixa edwardsii, driedfemale (TMCD363), Pingtung [preservedcoloration]; e, Ixa edwardsii, 

female (ZRC), Thailand; f, Myra biconica, driedfemale (TMCD419), Tainan [preserved 

coloration]; g, Nursia plicata, driedfemale (TMCD406), Anping [preserved coloration]; h, 

Pariphiculus mariannae, dried male (TMCD 436), Hengchun [preserved coloration].

Ng el al. : Checklist of crabs from Taiwan 

Fig. 3. a, Philyra heterograna, dried male (TMCD 427), Kaohsiung [preserved coloration]; b, 

Calappa bicornis, female (NTOU), Keelung; c, Calappa calappa, uniform coloured form, 

male (TMCD). Keelung; d, Calappa calappa, spotted form, male (TMCD), Keelung; e, 

Calappa capellonis, tlriedmale (TMCD 212), Taitung [preserved coloration] ]; f, Calappa 

capellonis, male (ZRC), Thailand; g, Mursia armata, male (ZRC 1999. 360), Nangfangao; h, 

Cyrtomaiacurviceros, male (NTOU), Tungsha. 

81

82 National Taiwan Museum Special Publication Series, No. 11. 86 pp.. April 200] 

Fig. 4. a* Platymaia bartschi, female (specimen in The Taiwan Fisheries Research Institute. 

Keelung), Tungsha (courtesyof D.-A. Lee); h, Platymaia fimbhata, female (ZRC), Tungsha 

(courtesy of D.-A. Lee); c, Pleistacantha sanctijohannis, dried female (TMCD455). Tailung 

[preservedcoloration]: d, Cyclocoelomatubervulata, driedmale (TMCD 1659), Kenling: e, 

Lissocarcinusarkati, driedfemale (TMCD480)„ Keelung fish market [preserved coloration!; 

f, Benthochasconhemingi, male (TMCD), Tungsha; g, h, Thalamitapelsartu male(NTOU). 

Kaohsiung.

Ng et al. : Checklist oi crabs from Taiwan 83 

Fig. 5. a, b. Scylla olivacea, male (ZRC), Tungkang; c, d, Scylla paramamosain, male (ZRC), 

Tungkang; e. f. Scylla serrata, male (ZRC), Tungkang; g, h, Scylla tranqueharica, male 

(ZRC), Tungkang.

National Taiwan Museum Special Publication Series, No. 11. 86 pp.. April 2001 

i#*_^fc- 

a, Demania cultripes, male (NTOU). Hopingtao, Keelung; b, Demania intermedia, female 

(NTOU), Hopingtao, Keelung; c, Demania japonica, male (ZRC), Tahsi; d, Demania reynaudi, 

male (NTOU), Hopingtao, Keelung; e, Demania rotundata, female (NTOU), Tahsi; f. 

Demania scaberrima, male (ZRC), Thailand; g, Demania toxica, male (NTOU), Nanhao. 

Kaohsiung; h, Juxtaxanthiaslividus, male (NTOU), Lanyu.


Fig. 7. a, Neoxanthops quadrilohatus, female (ZRC 1970. 3. 9. 2). Keelung [preserved coloration]; 

b, Actiteserythra, male (ZRC). Singapore; c, Carcinoplax indica, male {ZRC 1998. 216). 

Tahsi; d, Carcinoplax suruguensis, male, Nangfangao (courtesvofT. -Y. Chan): e, Carcinoplax 

tomentosa, male {ZRC 1998. 434), Nangfangao; f, Scalopidia spinosipes, male (ZRC), 

Thailand; g, Eucratealcocki, female (ZRC), Taiwan; h, EucrateSolaris, female (ZRC), 

Taiwan.

86 National Taiwan Museum Special Publication Series, No. 11. 86 pp.. April 2001 

* v ^ •?*•* 

Fig. 8. a, Chateau gramdatus, male (NTOU), Tahsi (courtesy of T.-Y. Chan); h. Macrophthalmus 

ceratophorus, female (NTOU), Tahsi; c, Macrophthalmus latreillel male (TMCD), Tainan; 

d, Ilyoplax delsmanu male (ZRC), Singapore; e, Pachygrapsus fakaravensis, male (ZRC), 

Hawaii; f, Helice fonnosensis, male (NTOU), Hsiangshan, Hsinchu; g, Helice subquadrata, 

female (NTOU), Hornet Taipei; h, Helice sp., male (NTOU), Lukang, Changhua

An annotated checklist of brachyuran crabs from Taiwan (Crustacea ...

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