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The Phylogeny, Systematics Goneplacidae Macleay (Crustacea

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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
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