The Phylogeny, Systematics Goneplacidae Macleay (Crustacea

Contributions to Zoology, 72 (2-3) 147-152 (2003)

SPB Academic Publishing bv, The Hague

The phylogeny, systematics and fossil record of the Goneplacidae MacLeay

(Crustacea, Decapoda, Brachyura) revisited

Hiroaki Karasawa¹ & Hisayoshi Kato²

Mizunami 2 1 Fossil Museum, Yamanouchi, Akeyo, Mizunami, Gifu 509-6132, Japan; Natural History

Museum and Institute, Chiba, Aoba-cho, Chiba 260-8682, Japan

Keywords:. Crustacea, Decapoda, Brachyura, Goneplacidae, phylogeny, systematics

Abstract based upon forty-five characters, and propose a new

classification (Appendix A) and phylogeny of the

We the and review the review Goneplacidae various alternative family. We suggest the division of the Goneplacidae hypotheses concerning membership within the family. We of- into six subfamilies, viz., Carinocarcinoidinae Kuro- fer a new cladistic based hypothesis ofphylogenetic relationships sawa & Kato, Chasmocarcininae, Euryplacinae, within the group. Goneplacinae (= Carcinoplacinae), Mathildellinae

Karasawa & Kato, and Trogloplacinae. Within the

Goneplacidae, the Trogloplacinae and Chasmocar- Introduction cininae are sister groups nested as the most derived

clade, followed by the Carinocarcinoidinae, Gone- Traditionally, the family Goneplacidae MacLeay placinae, Euryplacinae, and the most basal Ma- (Brachyura, Xanthoidea) has been recognized as a thildellinae. We also suggest the Pseudoziidae is monophyletic group (Balss, 1957). Since Guinot the sister group to the Eriphiidae. (1969a) first suggested that the Goneplacidae was

a polyphyletic group, the subfamilial arrangement has been modified by subsequent workers (Guinot, Results

1969b, 1971,1978; Manning & Holthuis, 1981; Ng,

1987 In and others). a recent systematic treatment, The Goneplacidae sensu lato has been commonly Lemaitre al. the Gone- et (2001) have now divided recorded from the Paleogene to the Recent and has placidae into six subfamilies, namely Carcinopla- included least previously at thirty-five fossil gen- cinae H. Milne Edwards, Chasmocarcininae Serene, era (Karasawa & Kato, in press). However, dis- Euryplacinae Pseudoziinae tinction Stimpson, Goneplacinae, between goneplacid on the one hand and Alcock, and Trogloplacinae Guinot. Subsequently, and panopeid, pilumnid, pseudorhombilid genera Bavie (2002) has two additional subfami- assigned on the other is difficult based solely upon carapace lies, Pilumnoidinae Guinot & Macpherson and Pla- characters (Schweitzer, 2000). A re-examination nopilumninae Serene, to the Goneplacidae, and of fossil taxa previously assigned to the Goneplaci- a t forded the Trogloplacinae full family status. Ng dae has shown that sixty-two species, twenty gen- & Liao excluded the Pseudoziinae from the (2002) and five subfamilies be era, may recognized as fossils Goneplacidae and elevated the Pseudoziinae to fa- (Karasawa & Kato, 2002, in press). Sixteen extinct 1111 ly status, and included the Planopilumninae and genera previously assigned to the family were not Pseudoziinae within the Pseudoziidae. referred to any goneplacid subfamilies and were In a recent we & in paper, (Karasawa Kato, press) excluded from the & , Goneplacidae (Karasawa Kato, Provide an adult morphology-based phylogenetic in press). In the same paper, we do not mention the a nalysis of fourteen within the genera Goneplacidae, systematic placement ofBicarinocarcinus Glaessner 148 H. Karasawa & H. Kato - Phytogeny of the Gonoplacidae

Fig. I. Strict consensus tree of four most-parsimonious trees oftwenty genera; phylogenetic analysis using PAUP* 4.0b (Swofford,

matrix in MacClade version Maddison, This tree is rooted 1999), data originating 4.05 (Maddison & 2002). against a hypothetical ancestor. Relative stability of clades was assessed using bootstrap (Felsenstein, 1985) and decay analyses (Bremer, 1994); bootstrap- ping was based on 100 replicates of random input order. The Bremer support was obtained using constraint trees generated by

Numbers branches and numbers branches MacClade and analyzed using PAUP*. above are bootstrap support below are Bremer

= support. Unambiguous character changes are as follows; box I = 45(1); 2 = 23(1), 24( 1); 3 27(1); 4 =36(1); 5 = 22(1), 25(1), 46(1);

6 =38(l),48(l); 7 = 15(1); 8 = 26(1), 28(1), 43(1); 9 = 16(1); 10 = 4(1); 11 = 1(0,37(1); 12 = 48(1); 13= 18(1), 19(1), 20(1), 21(1),

30(1), 32(1); 14 = 12(1), 17(1), 31(1); 15 = 13(1), 48(2).

& Secretan, 1987, which was originally placed with- domen fills the entire space between coxae ofpereio- in the The and thoracic the thoracic sternite does Carcinoplacinae. carapace pods 5, 8 not possess a

5 sternum characters are most like those of Carino- supplementary plate, dactyli of pereiopods are

sole sickle is twisted carcinoides Karasawa & Fudouji, the genus not shaped, gonopod 1 with a dis- of the Carinocarcinoidinae. Bicarinocar- tal and 2 is much shorter than Therefore, process, gonopod cinus is here referred to the Carinocarcinoidinae. gonopod 1. Thus, both genera lack the diagnostic

Schweitzer et al. (2002) have recently shown that characters of the Chasmocarcininae. Ng (2002)

Icriocarcinus there are close affinities between assigned Acidops Stimpson and Parapilumnus Koss-

Chasmocarcininae. Bishop and Ommatocarcinus White, and removed mann to the However, both ge- the former the Carcinerectidae Beurlen male abdomen that fills the entire genus from nera possess a and into the Goneplacidae. This occurrence extends space between coxae of pereiopods 5, have a nar- the thoracic with median sulcus the geologic range for the family back to the Late row sternum a on

Cretaceous. anterior part of sternite 4, an anterior margin of the

In more recent works, four genera have been male sterno-abdominal cavity which does not reach added to the Chasmocarcininae. Karasawa & Kato the anterior part of sternite 4, short pereiopods 2-

(in press) provisionally transfer Georgeoplax Tiirkay 5, and dactyli of pereiopods 2-5 terminating with and Litocheira Kinaham to the Pilumnidae, follow- acute chitinous tips. In Parapilumnus, the sulcus

6 ing Guinot (1969b, 1971), while Davie (2002) re- separating thoracic sternites and 7 is complete ferred both in genera to the Chasmocarcininae. They (Ng, 2002). These characters are not observed both differ from members of Chasmocarcininae members of the Chasmocarcininae (sensu Karasawa

(sensu & in we Karasawa Kato) that a wide male ab- & Kato) which is why exclude Acidops, Georgeo- Contributions to Zoology, 72 (2-3) - 2003 149

plax, Litocheira and Parapilumnus from the Chas- morphies (10-1, 45-1). D’Udekem d’Acoz (1999) mocarcininae. raised the Pilumnoidinaeto full family status. The

Karasawa & Kato (in do not discuss the the press) present analysis supports recognition of the subfamilial of placement Megaesthesius Rathbun, Pilumnoididae and suggests that the family is the

Notonyx A. Milne Raoulia and taxon Edwards, Ng, Typhlo- sister of the Carpiliidae. The monophyly of carcinodes Alcock, all ofwhich have been excluded the remaining goneplacids is well supported by four from the pilumnid subfamily Rhizopinae Stimpson synapomorphies (11 -1,22-1,25-1,46-1). However, by (1987). Serene (1964) Ng originally placed Mega- the present analysis is unable to resolve the rela- esthesius within his Chasmocar- new subfamily tionships betweenProgeryon (Goneplacidae incertae

cininae & Guinot while Davie (1996) and Karasawa sedis) and other goneplacid subfamilies.

& Kato excluded this from the Davie (in press) genus (2002) elevated the Trogloplacinae to full

subfamily. However, reas- Megaesthesius is here family status, although Davie & Guinot (1996) and

signed to the Chasmocarcininae based male Karasawa & upon Kato (in press) pointed out that the

abdomen and thoracic sternum characters. Serene Trogloplacinae has close affinities with the Chasmo- & Soh to the carcininae. If (1976) assigned Notonyx Goneplacinae the Trogloplacinae is treated as a

by a 2 a having long, elongate gonopod with long separate family, the remaining goneplacids become

flagellum; we concur. In Raoulia and a Typhlocar- polyphyletic group. D’Udekem d’Acoz (1999) cinodes, the male abdominal somites 3-5 are fused, raised the Euryplacinae and Carcinoplacinae to full

and the male 2 is and about to status and includedboth families gonopod long equal family in the super- I with gonopod a long flagellum. Therefore, both family ‘Goneplacoidea’. Stevcic (in Martin & Davis,

genera resemble members of the but also Trogloplacinae 2001) thought to elevate the Euryplacinae to detailed characters of the male thoracic sternum family status. There is a possibility that the six

are not known. subfamilies yet defined by Karasawa & Kato (in press) In the phylogenetic Karasawa& Kato be raised to full analysis by may family status, taking into ac- (in press), the Pilumnoidinae, assigned to the Gone- count of these works. placidae Davie (2002), was not included because It not by is clear which could be a reliable sister Guinot & (1987) noted there is a Macpherson that group to the Goneplacidae. Guinot (1969b) and close relationship between Pilumnoides Lucas and Stevcic (in Martin & Davis, 2001) mentioned that Carpilius Leach (Carpiliidae Ortmann), as based there is a close relationship between the Gone-

upon thoracic sternum and characters. Ng and based adult chelipcd placidae Geryonidae upon morphol- & Guinot transfer while (1999) suggested to Progeryon ogy, Rice (1980) showed that the family is Louvier from the Colosi the Geryonidae to Gone- most similar to the Pilumnidae Samouelle based

placidae. have re-examined adult Therefore, we an upon zoeal morphology. Von Sternberg & Cum- morphology-based for phylogenetic analysis twenty berlidge (2001) suggested that, based upon cladis- genera, including Carpilius Pilumnoides, andPro- tic and the , phenetic analyses, Goneplacidae might geryon, based 49 upon morphological characters be more closely related to the Portunoidea (inclu-

(Appendix B). Appendix C lists 49 characters and sive of the than of Geryonidae) to any family the character used in the states present analysis. Xanthoidea. Karasawa & Kato (in press) show that The present four the is derived analysis yielded most-parsimo- Goneplacidae as the sister group to ni °us trees, 108 steps long with a consistency in- the Pilumnidae. dex (Cl) of 0.6019, a retention index (RI) of 0.7962 The subfamilial arrangements of some genera a nd a rescaled index of consistency (RC) 0.4792. have not yet been satisfactorily cleared. Should the A strict of four consensus most-parsimonious trees, known subfamilies be given full family status? Then, 'ndicating bootstrap and Bremer is support, given should they be transferred from the superfamily 'n Fig. 1. the Pilumnoides, sole of the Pilum- Xanthoidea the What is , genus to ‘Goneplacoidea’? a true

n°idinae, is excluded from the Pilum- sister of the Goneplacidae. group Goneplacidae? These are sub- n°ides and Carpilius are sister taxa nested as the of jects forthcoming papers. 'Host basal clade and both share genera two synapo- 150 H. Karasawa & H. Kato - Phylogeny of the Gonoplacidae

Acknowledgements Lcmaitrc R, Gareia-Gomez J, von Sternberg R, Campos

NH. 2001. A of crab of new genus and a new species

family Goneplacidae MacLeay, 1838 (Crustacea: Decapoda: We thank P.K.L. Ng (National University of Singapore) for Brachyura) from the tropical western Pacific. Proc. biol. valuable comments on some goneplacidtaxa and on our phylo- Soc. Wash. 114: 951-963. genetic analysis, and D. Guinot (Museum national d’Histoire Maddison WP, Maddison DL. 2002. MacClade version 4.05. naturelle, Paris) and Z. Stevcic (Center for Marine Research, Massachusetts: Sinauer Associates. taxa. Croatia) for their much appreciated comments on some Manning RB, Holthuis LB. 1981. West African Brachyuran

crabs. Smiths. Contr. Zool. 306: 1-379.

Martin JW, Davis GB. 2001. An updated classification of

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Ng PKL, Guinot D. 1999. On a new species of deep-water 19(3B); xiv + 1-641. crab ofthe Progeryon (Decapoda, Brachyura, Geryo- Davie PJF, Guinot D. 1996. Two new freshwater crabs in genus nidae) from Hawai. Crustaceana 72: 685-692, Australocarcinus Davie, with remarks on Trogloplacinae Ng PKL. Liao LM. 2002. On a new species of Euryozius Guinot and Goneplacidae MacLeay (Crustacea: Decapoda: Miers, 1886 (Crustacea: Decapoda: Brachyura: Pseudo- Brachyura). Mem. Qld Mus. 39; 277-287. from the with ziidae) Philippines notes on the taxonomy D’Udckcm d’Acoz C. 1999. Invertaire et distribution des of the Proc. biol. Soc. Wash. 115: 585-593. crustaces decapodes de 1’Atlantique nord-oriental, de la genus.

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Karasawa Kato H. H, in press. The family Goneplacidae Received: 15 March 2003 1838 MacLeay, (Crustacea: Decapoda: Brachyura): sys-

tematics, phylogeny, and fossil records. Paleont. Res. 7. Contributions to Zoology, 72 (2-3) - 2003 151

Appendix A. Systematic list of to 10 fossae genera currently assigned the Antennular broad laterally: absent (0), present (1) Asterisks indicate extinct from Goneplacidae. genus (modified 11 Basal article ofantenna reaching front: present (0), absent Karasawa & Kato, in press). (1)

12 Ischium longer than merus: long (0), short (1)

Subfamily MathildellinaeKarasawa & in 13 Merus of Kato, press (Paleogene- maxilliped 3: subquadrate (0), suboval (1) Recent) 14 Telson about as long as wide (0), much longer than wide

Beuroisia Guinot & Richer de Forges, 1981,Branchioplax (1) Rathbun, 1916*,/ntes/i«Guinot&RicherdeForges, 1981, 15 Telson: triangular (0), suboval (1)

Mathildella Guinot& Richer de 1981, 16 Somites 4-6 much Forges, Neopilumno- narrower than 3: absent (0), present (1) plax Serene in 1969, Guinot, Platypilumnus Alcock, 1894, 17 Somite 3 much narrower than thoracic stemite 7: absent

1944* Tehuacana Stenzel, (0) present (1)

1871 Subfamily Euryplacinae Stimpson, (Eocene-Recent) 18 Somites 3 and 4: distinct (0), fused (1)

Chlinocephalus Ristori, 1886*, Corallicarcinus Muller & 19 Somites 4 and 5: distinct (0), fused (1)

Collins, 1991 *, Euryplax Stimpson, 1859,Eucrate de Haan, 20 Somites 3 and 4: movable (0), immovable (1) Fravillea Milne 1835, A. Edwards, 1880, Heteroplax Stimp- 21 Somites 4 and 5: movable (0), immovable (1): 1858, Machaerus Leach, 1818, Lemaitre 22 Sternum son, Nancyplax width: narrow (0), wide (1)

et al., 2001, Orbitoplax Tucker & Feldmann, 1990*,Psopheti- 23 Sulcus delimiting sternites 4 and 5: complete(0), interrupted

coides Sakai, 1969, Stoaplax Vega et al., 2001*, Trizo- medially (1)

carcinus Rathbun, 1914 24 Sulcus delimiting sternites 5 and 6: complete(0), interrupted

Viaplax Karasawa & Kato, in press* medially (1)

Subfamily Goneplacinae MacLeay, 1838 (Late Cretaceous- 25 Sulcus delimiting sternites 6 and 7: complete (0), interrupted

Recent): medially (I)

Bathyplax A. Milne 1880, H. Milne 26 Sulcus sternites Edwards, Carcinoplax delimiting 7 and 8: complete (0), interrupted Edwards, 1852, Goneplax Leach, 1814,Icriocarcinus Bis- medially (1)

hop, Neommatocarcinus Takeda & Median 1988*, Miyake, 1969, 27 sulcus on sternite 4: present (0), absent (1) A. MilneEdwards, Ommatocarcinus White, 28 Anterior end ofstemo-abdominal Notonyx 1873, cavity: posterior on sternite 1852, Psopheticus Wood-Mason, 1892,Singhaplax Serene 4 (0), anterior on 4 (1)

& Sob, 1976 29 of of Prolongation epistemite 7 male: absent (0), present

Subfamily Carinocarcinoidinae Karasawa & in Kato, press (Eo- (1) cene-Oligocene): 30 Stemite 7 laterally covered with sternite 8: absent (0), pre- Bicarinocarcinus Glaessner & Secretan, 1987*, Carino- sent (1)

carcinoides Karasawa & Fudouji, 2000* 31 Sternite 8 with supplementary plate; absent (0), present (1)

Subfamily Trogloplacinae Guinot, 1986 (Recent): 32 Sternite 8 visible ventrally; indistinct (0), distinct (1) Australocarcinus 1987, Guinot, 1986 33 Sternite visible Davie, Trogloplax 8 posteriorly: indistinct (0), distinct (1) Subfamily Chasmocarcininae Serene, 1964 (Eocene-Recent): 34 Gonopod 1: stout (0), slender (1)

Alcock & Chasmocarcinus Camatopsis Anderson, 1899, 3 5 Gonopod 1; slightly sinuous or curved (0), curved (1), sinuous Rathbun, 1898,Chasmocarcinops Alcock, 1900,Collinsius (2)

Gill- Karasawa, 1993*, Falconoplax van 1933*, 36 1 Straelen, Gonopod with hook-shaped apex: absent (0), present (1) carcinus Collins & Morris, 1978*, Hephthopelta Alcock, 37 Gonopod 1 with truncated apex: absent (0), present (1) 1899, Megaesthesius Rathbun, Bittner, 1884*, 38 1 inflated 1909, Mioplax Gonopod strongly proximally: absent (0), pre-

Orthakrolophos Schweitzer & Feldmann, 2001 *, Scalopidia sent (1)

Stimpson, 1858 39 Gonopod 2: long (0), short (1)

Goneplacidae incertae sedis 40 of 2: (Recent): Flagellum gonopod long (0), very short (1) Progeryon Bouvier, 1922, Raoulia Ng, 1987, 41 Gonopod 2 with wing-like flagellum: Typhlocar- absent (0), present cinodes Alcock, 1900 (!)

42 of Fingers pereiopods I elongate, much longer than palm:

absent (0), present (1) Appendix B. Characters and their states usedin PAUP* 43 analysis. Fingers of pereiopods 1 dark in color: present (0), absent (1) * Front with median notch: present absent (1) 44 of 1 with ventral absent (0), Carpus pereiopods spine: (0), pre- 2 Front with median projection: absent (0), present (1) sent (1) 2 Frontal teeth: absent present (0), (1) 45 Basis and ischium of pereiopods 1: distinct (0), indistinct 4 Notch between frontalmargin and supraorbital angle: distinct (1) (0), indistinct (1) 46 Meri of pereiopods 2-5 length: short (0), long (1) 2 Orbital moderate wide width: narrow (0), (1), (2) 47 of 2-5 with corneous Dactyli pereiopods tip: present (0), *2 Upper orbital fissure: present (0), absent (1) absent(1) 2 Dorsal more or less distinct region; (0), indistinct (1) 48 Dactyli ofpereiopods 5: styliform (0), spatulate (1), sickle- 8 Anterolateral teeth: >3 (0), 1-3 (1), 0 (2) shaped (2) 9 Eye stalk; short 49 of 5 with (0), long (1) Dactyli pereiopods setae: present (0), absent (1) 152 II. Karasawa & H. Kato - Phytogeny of the Gonoplacidae

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C. atocarcinus a Camatopsis Chasmocarcinus Trogloplax Carinocarcinoides Carcinoplax Goneplax Ommatocarcinus Psopheticus Heteroplax Mathildella Epixanthus Pseudozius Carpilius Pilumnoides Hypothetical Appendix TaxTaxa Goneplax Omni Eucrate Euryplax Heteroplax Beuroisia Intesius Progeryon Pilumnus Carpilius