ON THE TAXONOMY OF PSEUDOGRAPSUS SETOSUS (FABRICIUS, 1798) (DECAPODA, BRACHYURA, GRAPSIDAE

ON THE TAXONOMY OF PSEUDOGRAPSUS SETOSUS (FABRICIUS 1798)(DECAPODA BRACHYURA GRAPSIDAE)

BY

N K NG1 M-S JENG2 and PETER K L NG11 Department of Biological Sciences National University of Singapore Lower Kent Ridge Road

Singapore 119260 Republic of Singapore2 Institute of Zoology Academia Sinica Nankang Taipei 11529 Taiwan ROC

ABSTRACT

The taxonomy of Pseudograpsus setosus (Fabricius 1798) is discussed A neotype is designatedfor this species and a simultaneous lectotype is selected for two subjective junior synonyms CancerpenicilligerLatreille 1817 and Pseudograpsusbarbatus H Milne Edwards 1853 It is also recordedfrom Taiwan for the rst time and aspects of its ecology are discussed

REacuteSUMEacute

La taxonomie de Pseudograpsus setosus (Fabricius 1798) est discuteacutee Un neacuteotype est deacutesigneacutepour cette espegravece et un lectotype simultaneacute est seacutelectionneacute pour deux synonymes subjectifs juniorsCancer penicilliger Latreille 1817 et Pseudograpsus barbatus H Milne Edwards 1853 Lrsquoespegraveceest aussi signaleacutee de Taiwan pour la premiegravere fois et certains aspects de son eacutecologie sont discuteacutes

INTRODUCTION

The varunine genus Pseudograpsus H Milne Edwards 1837 (Grapsidae) cur-rently contains six species viz P albus Stimpson 1858 P crassus A Milne-Edwards 1868 P elongatus (A Milne-Edwards 1873) (= P erythraeus Kossman1877) P intermedius Chhapgar 1955 P nudus Stimpson 1858 and P setosus(Fabricius 1798) (= Alpheus setosus Weber 1795 (nomen nudum) = Grapsuspenicilliger Latreille 1817 = Pseudograpsus barbatus H Milne Edwards 1853)Most of these species are characterized by a relatively squarish carapace and theouter surface of the chela sometimes possessing a prominent tuft of setae (Tesch1917) Holthuis (1977) selected Cancer penicilliger Latreille 1817 as the typespecies for the genus a species which is now generally regarded as a junior syn-onym of Cancer setosus Fabricius 1798 The detailed taxonomy of Pseudograpsuscopy Koninklijke Brill NV Leiden 2002 Crustaceana 75 (6) 759-775Also available online wwwbrillnl

760 N K NG M-S JENG amp P K L NG

setosus (Fabricius 1798) is treated here The range of the species is extended toTaiwan and notes are provided of its ecology

Specimens examined are deposited in the Institute of Zoology AcademiaSinica (IZAS) Taipei Museacuteum national drsquoHistoire naturelle (MNHN) Paris Mu-seum Zoologicum Bogoriense (MZB) Bogor Naturhistorisches Museum (NMB)Basel Institute of Marine Biology National Taiwan Ocean University (NTOU)Keelung Rijksmuseum van Natuurlijke Historie (RMNH) Leiden SenckenbergMuseum (SMF) Frankfurt am Main Taiwan National Museum (TMCD) TaipeiUnited States National Museum (USNM) Smithsonian Institution WashingtonDC Zoological Museum of Amsterdam (ZMA) Amsterdam and ZoologicalReference Collection Raf es Museum of Biodiversity Research National Univer-sity of Singapore (ZRC) Singapore All measurements provided are of the cara-pace widths and lengths respectively The abbreviations G1 and G2 are used forthe male rst and second pleopods respectively

TAXONOMY

Family GRAPSIDAE MacLeay 1838

Subfamily VARUNINAE Alcock 1900

Pseudograpsus H Milne Edwards 1837

Pseudograpsus setosus (Fabricius 1798) ( gs 1-4)

ldquoCancer barbatusrdquo Rumphius 1705 26 pl 10 g 2Cancer setosus Fabricius 1798 339 Latreille 1803 372Grapsus penicilliger Latreille 1817b 431 Lamarck 1818 454 Desmarest 1825 130 pl 15 g 1

Latreille 1825 148Cancer penicilliger mdash Henschel 1833 204Pseudograpsus penicilliger mdash H Milne Edwards 1837 82 Lucas 1840 71 pl 3 g 3 Holthuis

1977 162Eriocheir penicilliger mdash De Haan 1835 31Pseudograpsus barbatus H Milne Edwards 1853 191 Heller 1865 52 De Man 1902a 99 103

De Man 1902b 506Pseudograpsus setosus mdash Kingsley 1880 205 De Man 1892 317 Tesch 1918 97 Holthuis

1959 116 Seregravene 1968 104 Ng amp Nakasone 1993 2

Material examined mdash Indonesia 1 male (neotype) (400 pound 353 mm) (RMNH-D-191) Amboncoll E W A Ludeking 1864 Other specimens 2 males (MZB-CRU-1448) Sungei (= river)Ilgamung Desa (= village) Persiapan Air Mancur Taman (= recreation park) Wisata Tuti AdagaeKec (= county) Perwakilan Alor Timur (= east) coll Sri Hartini 17 April 1997 2 females (MZB-CRU-1449) Desa (= village) Kamot Kec (= county) Perwakilan Alor Timur (= east) Pulau(= island) Alor coll Sri Hartini 17 April 1997 1 male (NMB-712a) river Toebah Ceram collL F de Beaufort 1920 1 female (RMNH-D-2550) river Wukur near Sikka Flores don et detJ G de Man 26 April 1930 11 males 2 females (SMF-5317) Ambon coll Hitubei Alang 1890

PSEUDOGRAPSUS SETOSUS (FABR) 761

2 males (SMF-365) Tubrina coll 1891 1 male 1 female (SMF-5317) Ambon no date 1 male(SMF-5325) Ternate det J G de Man 1902 2 males 2 females (ZMA) river near BombamgFlores coll M Weber 1889 2 males (ZMA) river Toebah west Ceram February 1910 mdash Asia ingeneral 1 male (ZMA) no other data 1 male (296pound230 mm) (dried damaged) (MNHN-B-13213)mers drsquoAsie coll Geiche mdash Taiwan 2 males (IZAS-72487) Geng-Fang river I-Lan county collK-X Lee 29 April 2000 1 male (NTOU) Tahsi I-Lan county coll K-X Lee September 20001 female (TMCD) Chang-Yuen Taitung county coll J-H Lee 28 August 2000 1 male 1 female(ZRC 20002230) Tahsi river I-Lan county Taiwan coll M-S Jeng 24 November 1997 2 males(ZRC) Gengfang river I-Lan county coll K-X Lee August 2000

Comparative material mdash P crassus A Milne-Edwards 1868 1 male (363pound313 mm) 1 female(364 pound 315 mm) (NHM-619a) Jomohon Celebes [= Sulawesi] coll 24 April 1894 1 male(426 pound 363 mm) (USNM 93152) Siaton river Siaton Municipality Province of Negros OrientalNegros Philippines coll D V Hart mdash P albus Stimpson 1858 2 males (98-106 pound 88-95 mm)2 females (80-92 pound 71-80 mm) (ZRC 20000529) Canton Island Hawaii coll O DegenerApril 1951 2 males (84-101 pound 78-90 mm) 2 females (ovigerous) (84-96 pound 73-85 mm) (ZRC20002231) Hou-Wan Kenting Pingtung county Taiwan coll H C Liu amp N K Ng 14 September1999

Diagnosis mdash Carapace quadrangular ( gs 1A 2A 3A) slightly longer thanbroad dorsal surface glabrous regions de ned convex Frontal margin slightlyconvex divided into four indistinct lobes Infraorbital margin nely granulatedinfraorbital region granulated ( g 3B) Anterolateral margin subcristate with threeteeth including the external orbital tooth external orbital tooth most distinct verybroad second tooth smaller in size third tooth smallest Posterolateral marginwith anterior part gently convex posterior part concave sharply converging toposterior carapace margin Orbits small eyes completely lling orbit corneaewell developed ( g 3B) Pterygostomial suborbital branchial regions granulatedhighly setose

Third maxilliped with foliaceous merus broader than long anterolateral anglestrongly produced auriculiform Ischium longer than broad sulcus not discernibleSmall distinct rhomboidal gape formed when third maxillipeds closed Exopodwith obtuse blunt inner subdistal angle agellum prominent longer than widthof merus Epistome broad at posterior margin granulated but without clefts or ssures ( g 3D)

Male chelae equal to subequal inner surfaces glabrous merus ( g 3E) withoutspines dorsal surface highly setose carpus without spines or teeth inner distalangle with broad low rounded lobe ( g 3F) outer surface of ngers with tuftsof long stiff black setae ( g 3G) inner surface with few setae at base of ngersFingers subequal to length of palm cutting edges of both ngers with numerousteeth nger tips recurved sharp hooved Female chelae ( g 3H) similar butsmaller than those of male without setae on dorsal surface of merus without longstiff setae on ngers ( g 3H)

Ambulatory legs with second pair longest ( gs 1A 2A) Merus of all ambu-latory legs with long soft setae anterior margin with blunt subdistal tooth outer


Fig 1 Pseudograpsus setosus (Fabricius 1798) male 385 pound 349 mm (ZRC 20002230) A dorsalview B anterior view C ventral view


Fig 2 Pseudograpsussetosus (Fabricius1798) female 355pound309 mm (ZRC 20002230)A dorsalview B anterior view C ventral view


Fig 3 Pseudograpsus setosus (Fabricius 1798) A-G I J male 385 pound 349 mm H female355 pound 309 mm (ZRC 20002230) A carapace B frontal view (setae not drawn) C thirdmaxilliped D epistome E dorsal view of merus of male chela F dorsal view of carpus of malechela G dorsal view of male cheliped H dorsal view of female cheliped I right third ambulatory

leg J right fourth ambulatory leg Scale A-J D 10 mm


Fig 4 Pseudograpsus setosus (Fabricius 1798) A C E F male 385 pound 349 mm B D G female355pound309 mm (ZRC 20002230)A male sternum B female sternum C male abdomen D female

abdomen E different views of G1 F different view of G2 G vulva Scale A-G D 10 mm

surface of carpus with short setae anterior and posterior margins of propodus withshort setae dactylus tapering to slender acute tip highly setose ( g 3I J)

Male abdomen narrowly triangular ( gs 1C 4C) lateral margins highly setose rst abdominal somite weakly arched with transverse ridge second somite


narrow short third somite broad slightly swollen laterally fourth somite broaderbut shorter than fth somite fth somite with proximal margin sinuous distalmargin with median part gently concave lateral margins gently convex sixthsomite quadrate lateral margins weakly convex proximal margin slightly concaveconvex medially telson broadly triangular lateral margins weakly concave distalmargin rounded Female abdomen very broad ( gs 2C 4D)

Lateral margins of rst 2 thoracic sternites nely granulated suture betweensternites 2 and 3 slightly convex towards abdomen lateral margins of sternites 3and 4 sinuous with deep broad notch demarcating edge of suture median groovebetween sternites 5 and 6 broad median part of sternite 8 longitudinally depressed( g 4A) Female thoracic sternum similar in form to male but relatively broader( g 4B)

Penis at outer edge of sternite 8 ( g 4A) G1 slender gently curving outwardsreaching to anterior margin of sternite 5 terminal lobe elongate slightly curvedgenital opening lateral to base of terminal lobe subterminal lobe pectinatedrounded densely setose ( g 4E) G2 short small ( g 4F) Vulva slightly convexcircular ( g 4G)

Colour mdash Live specimens are a uniform dark to chocolate brown preservedspecimens are of a lighter brown The setae on the male chelae are black even afterprolonged preservation in alcohol and drying

Remarks mdash This species now known as Pseudograpsus setosus (Fabricius1798) is generally regarded a senior subjective synonym of Grapsus penicilligerLatreille 1817 and Pseudograpsus barbatus H Milne Edwards 1853 The specieswas rst reported by Rumphius (1705 26 pl 10 no 2) as ldquoCancer barbatusrdquo onthe basis of specimen(s) from the Indonesian island of Ambon The descriptionwas very brief but a relatively good gure was provided which clearly showedthe long and stiff black setae at the base of the ngers of the male chela which isvery diagnostic of this taxon Rumphiusrsquo name ldquoCancer barbatusrdquo however hasno nomenclatural validity as it is a pre-Linnaean name Fabricius (1798 339) laterdescribed Cancer setosus on the basis of an unspeci ed number of specimens fromldquoTranquebariardquo (India) from the collections of Daldorff His description howeveris very brief and not very informative and he did not mention or refer to Rumphiusor his material

Latreille (1803 372) subsequently referred to a crab using the vernacular nameldquoCrabe Soyeuxrdquo (or lsquoSilk Crabrsquo) from Tranquebar and noted it was the Cancersetosus of Fabricius (1798) He did not indicate if he had actually examinedspecimens Subsequently Latreille (1817a 16 pl 12 g 1) identi ed and guredanother crab with reference to Rumphius (1705) but used another vernacular nameldquoGrapse Porte-Pinceaurdquo [or brush carrying crab] although he did not apply any


Latin name or suggest it was Cancer setosus Fabricius 1798 In another paperLatreille (1817b 431) used the Latin binomial name ldquograpsus penicilligerrdquo for hisldquoGrapse Porte-Pinceaurdquo and also referred to Rumphiusrsquo gure noting that it wasdistinctive in its possession of a strong tuft of black setae at the bases of its ngersHe did however indicate that he had seen at least one specimen in the collectionsof the Paris Museum Under current zoological nomenclatural rules Latreillersquos(1817b) use of ldquograpsus penicilligerrdquo is valid Henri Milne Edwards (1837)transferred Grapsus penicilliger Latreille 1817 to a new genus Pseudograpsustogether with a new species P pallipes However in a footnote H Milne Edwards(1837 82) questioned the validity of Grapsus penicilliger commenting that it wasprobably synonymous with Cancer setosus Fabricius 1798 From his text it is alsoclear that H Milne Edwards examined at least one specimen in the Paris Museum(probably the same one referred to in Latreille 1817b 431 see later) LaterH Milne Edwards (1853 191) formally synonymized Cancer setosus Fabricius1798 and Grapsus penicilliger Latreille 1817 under ldquoPseudograpsus barbatus(Rumphius 1705)rdquo However since ldquoCancer barbatus Rumphius 1705rdquo is not anavailable name (being pre-Linnaean) H Milne Edwards (1853) was actually the rst worker to validate the name ldquoPseudograpsus barbatusrdquo and the authorship ofthis name is his (see Tesch 1918) This makes Pseudograpsus barbatus H MilneEdwards 1853 a junior synonym of both Cancer setosus Fabricius 1798 andGrapsus penicilliger Latreille 1817

Grapsus penicilliger Latreille 1817 and Pseudograpsus barbatus H MilneEdwards 1853 both refer to Rumphiusrsquo (1705) ldquoCancer barbatusrdquo and Rumphiusrsquospecimens are thus syntypes Of course Rumphiusrsquo (1705) material is now lostThe specimen(s) of Grapsus penicilliger Latreille 1817 and Pseudograpsusbarbatus H Milne Edwards 1853 in the Paris Museum which were examined byLatreille (1817b) and H Milne Edwards (1837 1853) are also syntype(s) of theirrespective species With the help of Daniegravele Guinot the rst author managed to nda single dried damaged and very old male specimen in the Paris Museum (MHNHB-13213) labelled as Pseudograpsus setosus supposedly from the ldquoMers drsquoAsierdquoThere are no other specimens (preserved wet or dried) in the Paris MuseumAccording to the museumrsquos register this specimen had originally been listed asPseudograpsus barbatus There are no indications in the registrar or on the labelthat it is a type specimen or if it was the one examined by Latreille (1817b) andorH Milne Edwards (1853) but this is a common problem of that time types weresimply not formally designated The specimen could also have been obtained wellafter their time the original material being actually lost The available informationhowever strongly suggests that this dried male specimen (MNHN B-13213) wasthe same one used by Latreille (1817b) and H Milne Edwards (1853) and shouldbe regarded as a syntype of both species We hereby designate this specimen as


the lectotype of Grapsus penicilliger Latreille 1817 and Pseudograpsus barbatusH Milne Edwards 1853 an action that effectively makes both names objectivesynonyms

The type specimen of Cancer setosus Fabricius 1798 is no longer of anyscienti c use Zimsen (1964 650) notes that there are only dried remains of thespecimen labelled as this species in the Copenhagen Museum We have on hand aphotograph taken in the late 1960s of these remains which show only a few badlybroken-up pieces of legs These remains have since been rehydrated and the thirdauthor examined this material in late 1999 This material is in such poor conditionthat it is effectively useless and one cannot even discern the genus they belong toalthough all indications are that they probably belong to a varunine The type forCancer setosus should be regarded as no longer usable

In describing Pseudograpsus H Milne Edwards (1837 81) named two speciesas belonging to the genus P penicilliger (Latreille 1817) and P pallipes H MilneEdwards 1837 No type species was designated by him Holthuis (1977 162)was the rst to validly select Grapsus penicilliger Latreille 1817 as the typespecies (see also Ng amp Nakasone 1993 2) However Holthuis (1977) followingconvention also noted that Grapsus penicilliger Latreille 1817 is a juniorsynonym of Pseudograpsus setosus (Fabricius 1798)

Henri Milne Edwardsrsquo (1837 1853) belief that Cancer setosus Fabricius 1798is synonymous with Grapsus penicilliger Latreille 1817 (and Pseudograpsusbarbatus H Milne Edwards 1853) has been followed by almost all subsequentauthors without question But there are problems Fabricius (1798 339) haddescribed Cancer setosus very brie y ldquoC thorace utrinque bidentato tarsis hirtisHabitat Tranquebariae Dom Daldorff Corpus paruum laeuiusculum margineutrinque acuto bidentato Orbita oculorum haud eleuata Chelae breues inermespilosae Pedum tarsi hirtirdquo This description is too brief to ascertain its identityAlthough it notes that the chela [outer surface] was pilose presumably referringto the part near the base of the male ngers it does not describe the kindof setae present The presence of setae on the surface of the male chela is acharacter not unique to Pseudograpsus setosus or even members of the genusIt may also be present in several other varunine genera eg PtychognathusStimpson 1858 Hemigrapsus Dana 1851 Pyxidognathus A Milne-Edwards1879 or even Eriocheir De Haan 1835 On the basis of Fabriciusrsquo (1798) briefdescription Cancer setosus can be a species of Pseudograpsus Ptychognathus oreven Hemigrapsus The presence of long setae on the outer surface of the malechelae is the most prominent feature of P setosus and may have led to De Haanrsquosspeculative placement of this species in the genus Eriocheir (cf De Haan 183531) However it must be noted that whereas the setae present on the chelae of what


is now regarded as Pseudograpsus setosus are stiff and black those on Eriocheirspecies are soft and brown

On the other hand there is little doubt as to the identity of Grapsus penicilligerLatreille 1817 (= ldquoCancer barbatusrdquo Rumphius 1705 Pseudograpsus barbatusH Milne Edwards 1853) From Rumphiusrsquo (1705 pl 10 g 2) gure it is clearlythe species we are reporting from Taiwan ie a taxon with numerous long stiffblack setae at the base of its male ngers There is no other varunine species withsuch a character The setae retain their black colour even after long periods ofpreservation either in alcohol formalin or following drying Interestingly theIndonesian vernacular name of this species ldquoCattam Gigi Boeloerdquo (Rumphius1705 26) which means ldquocrab with hairy teethrdquo (alluding to the chelae) is stillused today by natives in Ambon and surrounding areas although their Englishname for it is ldquobearded crabrdquo (D Wowor pers comm)

Fabricius (1798 339) certainly does not mention the diagnostic stiff black setaepresent at the base of the male ngers of Grapsus penicilliger Latreille 1817Without this character one cannot be certain if Cancer setosus Fabricius 1798is actually the senior subjective synonym of Grapsus penicilliger Latreille 1817as is now generally accepted This is compounded by the fact that there are nocon rmed specimens of a varunine species from the Indian Ocean that has longstiff black setae on the outer surface of the male chela Since Fabriciusrsquo (1798)description of Cancer setosus from Tranquebar in southeastern India there hasonly been one other record of this species from the Indian Ocean ie by Heller(1865) from the Nicobar Islands (as Pseudograpsus barbatus) However whileHeller (1865) noted that his material (number and sex of specimens not stated)was characterized by the possession of a prominent tuft of setae at the base ofits cutting ngers he did not record whether these setae were black or stiffNevertheless on the basis of the synonymy he provided for the species Heller(1865) must be assumed to have been very aware of the papers of RumphiusLatreille and H Milne Edwards and he clearly followed H Milne Edwardsrsquo(1853) decision on the identities and names of the species concerned Thereare no other records of ldquoCancer setosusrdquo (as a Pseudograpsus or otherwise)from southeastern India Andaman or the Nicobar Islands (Chopra amp Das1937 Sethuramalingam amp Khan 1991 O F Fernando amp A F Fernandopers comm) Strangely Alcock (1900) made no mention of Cancer setosusGrapsus penicilliger or Pseudograpsus barbatus in his synopsis of the Indianfauna The rst author in March 2001 tried to obtain fresh specimens fromTranquebar and surrounding areas in Tamil Nadu State but this was not successfulNeither did the University Museum at Annamalai University which has an oldcollection possess any specimen Interviews with local biologists studying thefauna of Tranquebar revealed that they have not seen a crab species in which


the outer surface of the male chela has numerous long stiff black setae Onthe other hand they are familiar with Ptychognathus altimanus (Rathbun 1914)and Pseudograpsus intermedius Chhapgar 1955 (see also Sethuramalingam ampKhan 1991) both species possessing long brown soft setae at the base ofthe ngers of the male chela and they super cially resemble Cancer setosusand Grapsus penicilliger Ecological and feeding studies of the mangrove crabsin the Pitchavaram mangroves near Tranquebar have only found Ptychognathusaltimanus and Pseudograpsus intermedius (cf S Ravichandran pers comm) Thethird author has also studied several collections of freshwater decapods from theAndamans collected by Indraneil Das and his associates there are no specimensof what is now regarded as Pseudograpsus setosus In addition surveys of themarine fauna of the Andaman and Nicobar Islands have not uncovered any specieswith long black stiff setae on the chela (G Sivaleela Chennai Marine BiologicalStation Zoological Survey of India) It is also noteworthy that all the material ofwhat is now regarded as Pseudograpsus setosus from the Paci c has been collectedfrom islands Ambon Ceram Flores and Taiwan there are no known specimensfrom any mainland locality This casts further doubt on the identity of Fabriciusrsquo(1798) Cancer setosus from Tranquebar on mainland India although it is possiblethat Hellerrsquos (1865) record from the Nicobar Islands is actually correct It is alsoimportant to note that the area surrounding Tranquebar is relatively at with thecoastline lined with extensive sandy shores or mangroves This habitat is verydifferent from what we know of that preferred by Pseudograpsus setosus iesteep rocky mountain streams that enter the sea rapidly or if there is a sandybeach this stretches only a short distance from the shore

As such while the identities of Grapsus penicilliger Latreille 1817 andPseudograpsus barbatus H Milne Edwards 1853 are now clear with the above-mentioned designation of a lectotype the absence of a usable type-specimen forCancer setosus is far more serious especially since there are no clear records of aspecies with long stiff black setae on the male ngers from the Indian Ocean

The available data actually suggest that Cancer setosus Fabricius 1798 may infact not be synonymous with Grapsus penicilliger Latreille 1817 But this cannever be ascertained due a general lack of specimens from the western IndianOcean many of the islands there have not been well surveyed and what is nowregarded as Pseudograpsus setosus is after all a rather cryptic taxon (consider thesituation for this species in Taiwan where it was only discovered very recently)The absence of a usable type-specimen of Cancer setosus Fabricius 1798 makesthe resolution of this problem extremely dif cult There is at least a need tostabilize the name of the species from Taiwan as this is now the subject of severalongoing ecological and conservation studies by various workers there To usethe name Grapsus penicilliger Latreille 1817 for the Taiwan and Indonesian


populations is not satisfactory partly because it has not been used as a validname for over 150 years Also since H Milne Edwards (1853) Cancer setosusFabricius 1798 has been generally regarded as a senior synonym of Grapsuspenicilliger Latreille 1817 This of course does not help resolve the identityof Cancer setosus Fabricius 1798 as discussed earlier it can just as well beconspeci c with several other taxa (eg Ptychognathus altimanus (Rathbun 1914)or Pseudograpsus intermedius Chhapgar 1955) Under the present circumstancesit is clearly unsatisfactory to leave the matter of the identity of Cancer setosusFabricius 1798 in limbo

We believe the best solution is to select a neotype for Cancer setosus Fabricius1798 so as to retain the name Cancer setosus Fabricius 1798 for the species as isgenerally accepted today This is the only action that will not cause any confusionand will not result in any future taxonomic or nomenclatural uncertainties orproblems To this end a request will be sent to the International Commission onZoological Nomenclature to use its Plenary Powers to suppress the fragments in theCopenhagen Museum as the type of Cancer setosus and instead designate a malespecimen measuring 400 by 353 mm (RMNH-D-191) collected from Ambonas the neotype of Cancer setosus Fabricius 1798 With regards to the dried anddamaged specimen in the Paris Museum which is here regarded as the lectotypeof Grapsus penicilliger Latreille 1817 and Pseudograpsus barbatus H MilneEdwards 1853 its selection as the neotype would be highly unsatisfactory as itis in poor condition and lacks precise locality data The neotype selected at presentwas the specimen reported by Tesch (1918 97) This action keeps Cancer setosusFabricius 1798 as the subjective senior synonym of Grapsus penicilliger Latreille1817 and Pseudograpsus barbatus H Milne Edwards 1853 and thus maintains thestatus quo

Currently there is only one other species of Pseudograpsus known from TaiwanP albus Stimpson 1858 (see Suzuki 1985 Ng et al 2001) Pseudograpsussetosus however can easily be distinguished from P albus by the followingcharacters (a) presence of three distinct anterolateral teeth including the orbitaltooth (vs three anterolateral lobes) (b) presence of numerous tufts of long stiffsetae on the ngers of the chelae (vs a single tuft of short soft setae) and (c) agenerally larger adult size being mature at about 385 by 349 mm (vs specimensalready mature at about 98 by 73 mm [ZRC 20000529]) Pseudograpsus albusis also a wholly marine species unlike P setosus which regularly moves intofreshwater

Of its congeners P setosus is most similar to P crassus A Milne-Edwards1868 which is also a relatively larger-sized species Both species are also found infreshwater streams sometimes at high altitudes Pseudograpsus setosus has beenfound at altitudes of some 300 m P crassus up to 800 m (Tesch 1918) However


P setosus can be differentiated from P crassus by the following characters(a) the anterolateral margin is relatively less vaulted in P setosus (b) the thirdanterolateral tooth is relatively more acute (vs very low and relatively blunter)(c) the pterygostomial region is setose (vs glabrous) (d) the distal dorsal marginof the merus of the third maxlliped is relatively more foliose (e) presence of a rowof granules just above the moulting suture (vs absence of granules) (f) the dorsalsurface of the merus of the chelae is prominently setose (vs glabrous) (g) thechelae of males have tufts of long stiff setae on the outer surface of the pollex andmanus (vs glabrous) (h) the propodus of the ambulatory legs is densely fringedwith long soft setae (vs slightly fringed with short soft setae) (i) a relativelystouter merus of the last ambulatory leg with a length to width ratio of 32 (vs amore slender merus length to width ratio 43) (j) a stouter propodus of the lastambulatory leg with a length to width ratio of 17 (vs a more slender propoduslength to width ratio 20) (k) a relatively short and stout G1 (vs long and slender)and (l) absence of a long apical process on the G1 (vs presence of a long apicalprocess on the G1)

In any case the genus Pseudograpsus is in need of a revision Ng amp Nakasone(1993) discussed this brie y when they removed P laniger Tesch 1918 togetherwith a new species (S enodis) to a new genus Scutumara (see also Ng amp Komai1999) The genus is currently being reappraised by the rst author as a part of herrevision of the Varuninae

Pseudograpsus setosus was previously known from southeastern India NicobarIslands (Indian Ocean localities dubious as discussed earlier) as well as the islandsof Ambon Timor Flores and Ternate in Indonesia (Fabricius 1798 Heller 1865De Man 1892 Tesch 1918 Holthuis 1978) The present discovery of P setosusfrom Taiwan is very interesting as this extends its known range considerably andinto northern temperate waters for the rst time In Taiwan it is known from thenortheastern and eastern parts of the island but there have been reliable reportsthat it also occurs in the southeast (P-H Ho pers comm) All known recordsare however from eastern localities its distribution being similar to the betterknown varunine Platyeriocheir formosa Chan Hung amp Yu 1998 (see Ng et al1999)

The biology of this species is not clearly known Most of the present specimenswere collected in the lower stream near the mouth of the Tachi and Genfang riversI-Lan county Taiwan However it has been observed that this species is present inthe moutains at about 300 m above sea level Migrating downstream to the rivermouth in the months of March and April and in the months of June and July theyhave been found in the sea itself being caught in shermanrsquos nets (pers commK-X Lee)


ACKNOWLEDGEMENTS

The authors would like to express their deepest gratitude to the referees whohave read through the manuscript laboriously and provided precious commentsAlso to the following curators of the various museums for their hospitality andhelp when the rst author was examining specimens in their respective institutesProf D Guinot (MNHN) Dr C H J M Fransen (RMNH) Mr D Platvoet(ZMA) Dr A Haeggi (NHM) and Dr M Tuumlrkay (SNG) Thanks are due toDr R Lemaitre (USNM) and Dr A Haeggi (NMB) for the loan of the P crassusspecimens to Miss D Wowor (MZB) for the loan of P setosus specimensand for her help in the translation of the Indonesian labels Dr C-H Wang(TMCD) and Dr H-P Ho (National Museum of Marine Biology and AquariumTaiwan) for their help in providing additional information on the habits andbiology of this species Mr Li Kuan-Xin (Peikuan Crab Museum Taiwan)kindly passed-on specimens to us and shared his knowledge of the species withthe authors Professor Lipke Holthuis helped check and correct the manuscriptmaking many important suggestions Thanks are due to Prof A F FernandoDr O F Fernando (Centre of Advanced Study in Marine Sciences AnnamalaiUniversity) Dr S Ravichandran (ENVIS Centre Annamalai University) andMs G Sivaleela (Marine Biological Station Zoological Survey of India) forproviding additional information about the crab fauna of Tranquebar Parangipettai(Porto Novo) and the Andaman and Nicobar Islands respectively Dr IndraneilDas kindly sent me several lots of crabs from the Andamans The authors wouldalso like to thank Lady McNiece Yuen Ping for her nancial support enabling the rst author to visit the various European museums to study the collections Theauthors would also like to thank Mr Yip Hoi Kee for developing the photographicprints Support from research grant R-154-00-044-113 to the third author from theNational University of Singapore is acknowledged This is a contribution from theEcology and Systematics Laboratory Department of Biological Sciences NationalUniversity of Singapore

REFERENCES

ALCOCK A 1900 Materials for a carcinologicalfauna of India No 6 The Brachyura Catometopaor Grapsoidea Journ Asiatic Soc Bengal 69 (2) 279-456

CHHAPGAR B F 1955 On two new species and a new variety of crabs (Decapoda Brachyura)from Bombay State Rec Indian Mus 53 (1 2) 251-260

CHOPRA B amp K N DAS 1937 Further notes on the Crustacea Decapoda in the Indian MuseumIX On three collections of crabs from Tavoy and Mergui Archipelago Rec Indian Mus 39423-434 pl 6

DESMAREST A G 1825 Consideacuteration geacuteneacuterales sur la classe des Crustaceacutes et description desespegraveces de ces animaux qui vivent dans la mer sur les cocirctes ou dans les eaux douces de laFrance 1-446 (Le Normant Fils imprimeur du Roi Paris)


FABRICIUS J C 1798 Supplementum Entomologiae systematicae 1-573 (Proft et StorchHafniae)

HAAN W DE 1833-1850 Crustacea In P F VON SIEBOLD (ed) Fauna Japonica sive descriptioanimalium quae in itinere per Japoniam jussu et auspiciis superiorum qui summum inIndia Batava Imperium tenent suscepto annis 1823-1830 collegit notis Observationibus etadumbrationibus illustravit 1-243 (Lugduni-Batavorum)

HELLER C 1865 Crustacea Reise der oesterreichischen Fregatte Novara um die Erde in denJahren 1857-1858-1859unter den Befehlen des Commodors B von Wuumlllerstorf-Urbair(Zool)2 (3) 1-280 pls 1-25 (Vienna)

HENSCHEL A G E T 1833 Vita G E Rumphii plinii indici Accedunt specimen materiaeRumphianae medicae clavisque herbarii et thesauri Amboinensis i-xiv 1-215 1 pl

HOLTHUIS L B 1959 Notes on pre-linnean carcinology (including the study of Xiphosura) ofthe Malay Archipelago In H C D DE W IT Rumphius memorial volume 63-125 7 pls(Hollandia NV Baarn)

mdash mdash 1977 The Grapsidae Gecarcinidae and Palicidae (Crustacea Decapoda Brachyura) of theRed Sea Israel Journ Zool 26 141-192

mdash mdash 1978 A collectionof decapod Crustacea from Sumba Lesser Sunda Islands Indonesia ZoolVerh Leiden 162 1-55

KINGSLEY J S 1880 Carcinological notes No 4 Synopsis of the Grapsidae Proc Acad natSci Philadelphia 1880 187-224

KOSSMANN R 1877 Malacostraca In ZoologischeErgebnisse einer in Auftrage der KoumlniglichenAcademie der Wissenschaften zu Berlin ausgefuumlhrten Reise in die Kuumlstengebiete des RothenMeeres 1 (3) (1) 1-66 pls 1-3

LAMARCK J B P A (MONET CHEVALIER DE) 1818 Histoire naturelle des animaux sansverteacutebreacutes preacutesentant les caractegraveres geacuteneacuteraux et particuliers de ces animaux leur distributionleurs classes leurs familles leurs genres et la citation des principales espegraveces qui srsquoyrapportent preacuteceacutedeacutee drsquoune introduction offrant la deacutetermination des caractegraveres essentielsde lrsquoanimal sa distinction du veacutegeacutetal et des autres corps naturelles en n lrsquoexposition desprincipes fondamentaux de la zoologie 5 1-612

LATREILLE P A 1802 Histoire naturelle geacuteneacuterale et particuliegravere des Crustaceacutes et des Insectes3 1-468 (F Dufort Paris)

mdash mdash 1817a Les Crustaceacutes les Arachnides et les Insectes In G CUVIER (ed) Le Regravegne Animal3 i-xxix 1-653 (A Belin Paris)

mdash mdash 1817b Grapsus Nouv Dict Hist nat (ed 2) 13 431mdash mdash 1825 Plagusie Plagusia Encycl meacuteth Hist nat (Ins) 10 145-148mdash mdash 1829 Les Crustaceacutes les Arachnides et les Insectes distribueacutes en families naturellesOuvrage

formant les tomes 4 et 5 de celui de M le Baron Cuvier sur Le Regravegne Animal 1-584 (Paris)LUCAS H 1840 Histoire naturelle des Crustaceacutes des Arachnides et des Myriapodes Ouvrage

accompagneacute de 46 planches graveacutees sur agier repreacutesentant plus de 300 sujets (ed 1) 1-600(P Dumeacuteril Paris)

MACLEAY W S 1838 Illustrationsof the Annulosa of South Africa being a portion of the objectsof natural history collected during an expedition into the interior of South Africa under thedirection of Dr Andrew Smith in the years 1834 1835 and 1836 tted out by ldquoThe Capeof Good Hope Association for Exploring Central Africardquo In A SMITH Illustrations of thezoology of South Africa consisting chie y of gures and descriptions of the objects of naturalhistory collected during an expedition into the interior of South Africa in the years 1834 1835and 1836 tted out by ldquoThe Cape of Good Hope Association for Exploring Central Africardquo(Invertebratae)1-75 4 pls (London)

MAN J G DE 1892 Decapoden des Indischen Archipels In M WEBER (ed) ZoologischeErgebnisse einer Reise in Niederlaumlndisch Ost-Indien 2 18-722 pl 17


mdash mdash 1902a Over de Crustacea (weeke schaalvisschen) in lsquoRumphiusrsquo Rariteitkamer RumphiusGedenkboek 98-104

mdash mdash 1902b Die von Herrn Professor Kuumlkenthal im Indischen Archipel gesammelten Dekapodenund Stomatopoden Abh Senckenberg naturf Gesellsch 25 467-929 pls 10-27

MILNE-EDWARDS A 1868 Description de quelques Crustaceacutes nouveaux provenant des voyagesde M Alfred Grandidier agrave Zanzibar et agrave Madagascar Nouv Arch Mus Hist nat Paris 469-92 pls 19-21

mdash mdash 1873 Recherches sur la faune carcinologiquede la Nouvelle-Caleacutedonie 2 Nouv Arch MusHist nat Paris 9 155-332 pls 4-18

MILNE EDWARDS H 1837 Histoire naturelle des Crustaceacutes comprenant lrsquoanatomie la physiolo-gie et la classi cation de ces animaux 2 1-531 atlas 1-32 pls 1-42 (Paris)

mdash mdash 1853 Meacutemoire sur la famille des Ocypodiens suite Ann Sci nat Paris (Zool) 20 (3)163-228 pls 6-11

NG N K amp T KOMAI 2000 On the male of Scutumara enodis Ng amp Nakasone 1993 (CrustaceaDecapoda Brachyura Grapsidae) Proc biol Soc Washington 113 (1) 48-53

NG N K J GUO amp P K L NG 1999 On the generic af nities of Eriocheir leptognathusRathbun 1913 and Eriocheir formosa Chan Hung and Yu 1995 with description of a newgenus (Crustacea Grapsidae Varuninae) Journ Crust Biol 19 (1) 154-170

NG P K L amp Y NAKASONE 1993 Scutumara enodis a new genus and species of grapsid crab(Decapoda Grapsidae) from Okinawa Ryukyus Japan Crust Res Tokyo 12 1-6

NG P K L C-H WANG P-H HO amp H-T SHIH 2001 An annotated checklist of brachyurancrabs from Taiwan (Crustacea Decapoda) Natn Taiwan Mus spec Publn Ser Taipei 111-86 8 col pls

RATHBUN M J 1914 New species of crabs of the families Grapsidae and Ocypodidae [Scienti cresults of the Philippines cruise of the sheries Steamer lsquoAlbatrossrsquo 1907-1910 No 31] ProcU S natn Mus 47 69-85

RUMPHIUS G E 1705 DrsquoAmboinsche rariteitkamer behelzende eene beschryvinge van aller-hande zoo weeke als harde schaalvischen te weeten raare krabben kreeften en diergelykezeedieren als mede allerhande hoorntjes en schulpen die men in drsquoAmboinsche zee vindtdaar beneven zommige mineraalen gesteenten en soorten van aarde die in drsquoAmboinsche enzommige omleggende eilanden gevonden worden 1-340 60 pls (Amsterdam)

SEREgraveNE R 1968 The Brachyura of the Indo-West Paci c region In Prodromus for a check list ofthe (non-planctonic)marine fauna of South East Asia Spec Publn Singapore natn Acad SciNo 1 Fauna IIICc 3 33-112

SETHURAMALINGAM S amp S AJMAL KHAN 1991 Brachyuran crabs of Parangipettai Coast1-93 28 pls (Annamalai University Parangipettai)

STIMPSON W 1858 Prodromus descriptionis animalium evertebratorum quae in Expeditione adOceanum Paci cum Septentrionalem a Republica Federata missa Cadwaladaro Ringgold etJohanne Rodgers Ducibus observavit et descripsit pt 5 Crustacea Ocypodoidea Proc Acadnat Sci Philadelphia 10 93-111

SUZUKI H 1985 A list of the macro-crustaceans collected from Formosa Report Ocean EnvnSouthern Ryukyu Is 2 49-59

TESCH J J 1918 The Decapoda Brachyura of the Siboga Expedition I HymenosomidaeRetroplumidae Ocypodidae Grapsidae and Gecarcinidae Siboga-Exp Mon 39 (c2) 1-148pls 1-6

WEBER F 1795 Nomenclator entomologicus secundum Entomologiam systematicam ill Fabriciiadjectis speciebus recens detectis et varietatibus i-viii 1-171 (Chilonii amp Hamburgi)

ZIMSEN E 1964 The type material of J C Fabricius 1-656 2 pls (Copenhagen)

First received 14 March 2001Final version accepted 14 February 2002


setosus (Fabricius 1798) is treated here The range of the species is extended toTaiwan and notes are provided of its ecology

Specimens examined are deposited in the Institute of Zoology AcademiaSinica (IZAS) Taipei Museacuteum national drsquoHistoire naturelle (MNHN) Paris Mu-seum Zoologicum Bogoriense (MZB) Bogor Naturhistorisches Museum (NMB)Basel Institute of Marine Biology National Taiwan Ocean University (NTOU)Keelung Rijksmuseum van Natuurlijke Historie (RMNH) Leiden SenckenbergMuseum (SMF) Frankfurt am Main Taiwan National Museum (TMCD) TaipeiUnited States National Museum (USNM) Smithsonian Institution WashingtonDC Zoological Museum of Amsterdam (ZMA) Amsterdam and ZoologicalReference Collection Raf es Museum of Biodiversity Research National Univer-sity of Singapore (ZRC) Singapore All measurements provided are of the cara-pace widths and lengths respectively The abbreviations G1 and G2 are used forthe male rst and second pleopods respectively

TAXONOMY

Family GRAPSIDAE MacLeay 1838

Subfamily VARUNINAE Alcock 1900

Pseudograpsus H Milne Edwards 1837

Pseudograpsus setosus (Fabricius 1798) ( gs 1-4)

ldquoCancer barbatusrdquo Rumphius 1705 26 pl 10 g 2Cancer setosus Fabricius 1798 339 Latreille 1803 372Grapsus penicilliger Latreille 1817b 431 Lamarck 1818 454 Desmarest 1825 130 pl 15 g 1

Latreille 1825 148Cancer penicilliger mdash Henschel 1833 204Pseudograpsus penicilliger mdash H Milne Edwards 1837 82 Lucas 1840 71 pl 3 g 3 Holthuis

1977 162Eriocheir penicilliger mdash De Haan 1835 31Pseudograpsus barbatus H Milne Edwards 1853 191 Heller 1865 52 De Man 1902a 99 103

De Man 1902b 506Pseudograpsus setosus mdash Kingsley 1880 205 De Man 1892 317 Tesch 1918 97 Holthuis

1959 116 Seregravene 1968 104 Ng amp Nakasone 1993 2

Material examined mdash Indonesia 1 male (neotype) (400 pound 353 mm) (RMNH-D-191) Amboncoll E W A Ludeking 1864 Other specimens 2 males (MZB-CRU-1448) Sungei (= river)Ilgamung Desa (= village) Persiapan Air Mancur Taman (= recreation park) Wisata Tuti AdagaeKec (= county) Perwakilan Alor Timur (= east) coll Sri Hartini 17 April 1997 2 females (MZB-CRU-1449) Desa (= village) Kamot Kec (= county) Perwakilan Alor Timur (= east) Pulau(= island) Alor coll Sri Hartini 17 April 1997 1 male (NMB-712a) river Toebah Ceram collL F de Beaufort 1920 1 female (RMNH-D-2550) river Wukur near Sikka Flores don et detJ G de Man 26 April 1930 11 males 2 females (SMF-5317) Ambon coll Hitubei Alang 1890






















































ACKNOWLEDGEMENTS


REFERENCES

































































































ACKNOWLEDGEMENTS


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ACKNOWLEDGEMENTS


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ACKNOWLEDGEMENTS


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ACKNOWLEDGEMENTS


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ACKNOWLEDGEMENTS


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ACKNOWLEDGEMENTS


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ACKNOWLEDGEMENTS


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ACKNOWLEDGEMENTS


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ACKNOWLEDGEMENTS


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ACKNOWLEDGEMENTS


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ACKNOWLEDGEMENTS


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ACKNOWLEDGEMENTS


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ACKNOWLEDGEMENTS


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ON THE TAXONOMY OF PSEUDOGRAPSUS SETOSUS (FABRICIUS, 1798) (DECAPODA, BRACHYURA, GRAPSIDAE

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