Contributions to Zoology, 72 (2/3) (2003)
The phylogeny, systematics and fossil record of the Goneplacidae MacLeay (Crustacea, Decapoda, Brachyura) revisited
Hiroaki Karasawa , Hisayoshi Kato
Keywords: Crustacea, Decapoda, Brachyura, Goneplacidae, phylogeny, systematics
We review the Gonoplacidae and review the various alternative hypotheses concerning membership within the family. We offer a new cladistic based hypothesis of phylogenetic relationships within the group.
Traditionally, the family Goneplacidae MacLeay (Brachyura, Xanthoidea) has been recognized as a monophyletic group (Balss, 1957). Since Guinot (1969a) first suggested that the Goneplacidae was a polyphyletic group, the subfamilial arrangement has been modified by subsequent workers (Guinot, 1969b, 1971, 1978; Manning & Holthuis, 1981; Ng, 1987 and others). In a recent systematic treatment, Lemaitre et al. (2001) have now divided the Goneplacidae into six subfamilies, namely Carcinoplacinae H. Milne Edwards, Chasmocarcininae Serène, Euryplacinae Stimpson, Goneplacinae, Pseudoziinae Alcock, and Trogloplacinae Guinot. Subsequently, Davie (2002) has assigned two additional subfamilies, Pilumnoidinae Guinot & Macpherson and Planopilumninae Serène, to the Goneplacidae, and afforded the Trogloplacinae full family status. Ng & Liao (2002) excluded the Pseudoziinae from the Goneplacidae and elevated the Pseudoziinae to family status, and included the Planopilumninae and Pseudoziinae within the Pseudoziidae.
In a recent paper, we (Karasawa & Kato, in press) provide an adult morphology-based phylogenetic analysis of fourteen genera within the Goneplacidae, based upon forty-five characters, and propose a new classification (Appendix A) and phylogeny of the family. We suggest the division of the Goneplacidae into six subfamilies, viz., Carinocarcinoidinae Karasawa & Kato, Chasmocarcininae, Euryplacinae, Goneplacinae (= Carcinoplacinae), Mathildellinae Karasawa & Kato, and Trogloplacinae. Within the Goneplacidae, the Trogloplacinae and Chasmocarcininae are sister groups nested as the most derived clade, followed by the Carinocarcinoidinae, Goneplacinae, Euryplacinae, and the most basal Mathildellinae. We also suggest the Pseudoziidae is the sister group to the Eriphiidae.
The Goneplacidae sensu lato has been commonly recorded from the Paleogene to the Recent and has previously included at least thirty-five fossil genera (Karasawa & Kato, in press). However, distinction between goneplacid on the one hand and panopeid, pilumnid, and pseudorhombilid genera on the other is difficult based solely upon carapace characters (Schweitzer, 2000). A re-examination of fossil taxa previously assigned to the Goneplacidae has shown that sixty-two species, twenty genera, and five subfamilies may be recognized as fossils (Karasawa & Kato, 2002, in press). Sixteen extinct genera previously assigned to the family were not referred to any goneplacid subfamilies and were excluded from the Goneplacidae (Karasawa & Kato, in press). In the same paper, we do not mention the systematic placement of Bicarinocarcinus Glaessner & Secretan, 1987, which was originally placed within the Carcinoplacinae. The carapace and thoracic sternum characters are most like those of Carinocarcinoides Karasawa & Fudouji, the sole genus of the Carinocarcinoidinae. Therefore, Bicarinocarcinus is here referred to the Carinocarcinoidinae. Schweitzer et al. (2002) have recently shown that there are close affinities between Icriocarcinus Bishop and Ommatocarcinus White, and removed the former genus from the Carcinerectidae Beurlen and into the Goneplacidae. This occurrence extends the geologic range for the family back to the Late Cretaceous.
In more recent works, four genera have been added to the Chasmocarcininae. Karasawa & Kato (in press) provisionally transfer Georgeoplax Türkay and Litocheira Kinaham to the Pilumnidae, following Guinot (1969b, 1971), while Davie (2002) referred both genera to the Chasmocarcininae. They both differ from members of Chasmocarcininae (sensu Karasawa & Kato) in that a wide male abdomen fills the entire space between coxae of pereiopods 5, the thoracic sternite 8 does not possess a supplementary plate, dactyli of pereiopods 5 are not sickle shaped, gonopod 1 is twisted with a distal process, and gonopod 2 is much shorter than gonopod 1. Thus, both genera lack the diagnostic characters of the Chasmocarcininae. Ng (2002) assigned Acidops Stimpson and Parapilumnus Kossmann to the Chasmocarcininae. However, both genera possess a male abdomen that fills the entire space between coxae of pereiopods 5, have a narrow thoracic sternum with a median sulcus on the anterior part of sternite 4, an anterior margin of the male sterno-abdominal cavity which does not reach the anterior part of sternite 4, short pereiopods 2-5, and dactyli of pereiopods 2-5 terminating with acute chitinous tips. In Parapilumnus, the sulcus separating thoracic sternites 6 and 7 is complete (Ng, 2002). These characters are not observed in members of the Chasmocarcininae (sensu Karasawa & Kato) which is why we exclude Acidops, Georgeoplax, Litocheira and Parapilumnus from the Chasmocarcininae.
Karasawa & Kato (in press) do not discuss the subfamilial placement of Megaesthesius Rathbun, Notonyx A. Milne Edwards, Raoulia Ng, and Typhlocarcinodes Alcock, all of which have been excluded from the pilumnid subfamily Rhizopinae Stimpson by Ng (1987). Serène (1964) originally placed Megaesthesius within his new subfamily Chasmocarcininae while Davie & Guinot (1996) and Karasawa & Kato (in press) excluded this genus from the subfamily. However, Megaesthesius is here reassigned to the Chasmocarcininae based upon male abdomen and thoracic sternum characters. Serène & Soh (1976) assigned Notonyx to the Goneplacinae by having a long, elongate gonopod 2 with a long flagellum; we concur. In Raoulia and Typhlocarcinodes, the male abdominal somites 3-5 are fused, and the male gonopod 2 is long and about equal to gonopod 1 with a long flagellum. Therefore, both genera resemble members of the Trogloplacinae but detailed characters of the male thoracic sternum are not yet known.
In the phylogenetic analysis by Karasawa & Kato (in press), the Pilumnoidinae, assigned to the Goneplacidae by Davie (2002), was not included because Guinot & Macpherson (1987) noted that there is a close relationship between Pilumnoides Lucas and Carpilius Leach (Carpiliidae Ortmann), as based upon thoracic sternum and cheliped characters. Ng & Guinot (1999) suggested to transfer Progeryon Bouvier from the Geryonidae Colosi to the Goneplacidae. Therefore, we have re-examined an adult morphology-based phylogenetic analysis for twenty genera, including Carpilius, Pilumnoides, and Progeryon, based upon 49 morphological characters (Appendix B). Appendix C lists 49 characters and character states used in the present analysis.
The present analysis yielded four most-parsimonious trees, 108 steps long with a consistency index (CI) of 0.6019, a retention index (RI) of 0.7962 and a rescaled consistency index (RC) of 0.4792. A strict consensus of four most-parsimonious trees, indicating bootstrap and Bremer support, is given in Fig. 1. Pilumnoides, the sole genus of the Pilumnoidinae, is excluded from the Goneplacidae. Pilumnoides and Carpilius are sister taxa nested as the most basal clade and both genera share two synapomorphies (10-1, 45-1). D’Udekem d’Acoz (1999) raised the Pilumnoidinae to full family status. The present analysis supports the recognition of the Pilumnoididae and suggests that the family is the sister taxon of the Carpiliidae. The monophyly of the remaining goneplacids is well supported by four synapomorphies (11-1, 22-1, 25-1, 46-1). However, the present analysis is unable to resolve the relationships between Progeryon (Goneplacidae incertae sedis) and other goneplacid subfamilies.
Fig. 1. Strict consensus tree of four most-parsimonious trees of twenty genera; phylogenetic analysis using PAUP* 4.0b (Swofford, 1999), data matrix originating in MacClade version 4.05 (Maddison & Maddison, 2002). This tree is rooted against a hypothetical ancestor. Relative stability of clades was assessed using bootstrap (Felsenstein, 1985) and decay analyses (Bremer, 1994); bootstrapping was based on 100 replicates of random input order. The Bremer support was obtained using constraint trees generated by MacClade and analyzed using PAUP*. Numbers above branches are bootstrap support and numbers below branches are Bremer support. Unambiguous character changes are as follows: box 1 = 45(1); 2 = 23(1), 24(1); 3 = 27(1); 4 = 36(1); 5 = 22(1), 25(1), 46(1); 6 = 38(1), 48(1); 7 = 15(1); 8 = 26(1), 28(1), 43(1); 9 = 16(1); 10 = 4(1); 11 = 1(1), 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).
Davie (2002) elevated the Trogloplacinae to full family status, although Davie & Guinot (1996) and Karasawa & Kato (in press) pointed out that the Trogloplacinae has close affinities with the Chasmocarcininae. If the Trogloplacinae is treated as a separate family, the remaining goneplacids become a polyphyletic group. D’Udekem d’Acoz (1999) raised the Euryplacinae and Carcinoplacinae to full family status and included both families in the superfamily ‘Goneplacoidea’. Stevcic (in Martin & Davis, 2001) also thought to elevate the Euryplacinae to family status. There is a possibility that the six subfamilies defined by Karasawa & Kato (in press) may be raised to full family status, taking into account of these works.
It is not clear which could be a reliable sister group to the Goneplacidae. Guinot (1969b) and Stevcic (in Martin & Davis, 2001) mentioned that there is a close relationship between the Goneplacidae and Geryonidae based upon adult morphology, while Rice (1980) showed that the family is most similar to the Pilumnidae Samouelle based upon zoeal morphology. Von Sternberg & Cumberlidge (2001) suggested that, based upon cladistic and phenetic analyses, the Goneplacidae might be more closely related to the Portunoidea (inclusive of the Geryonidae) than to any family of the Xanthoidea. Karasawa & Kato (in press) show that the Goneplacidae is derived as the sister group to the Pilumnidae.
The subfamilial arrangements of some genera have not yet been satisfactorily cleared. Should the known subfamilies be given full family status? Then, should they be transferred from the superfamily Xanthoidea to the ‘Goneplacoidea’? What is a true sister group of the Goneplacidae? These are subjects of forthcoming papers.
Balss H. 1957. Decapoda. In: Bronn HG (ed.). Klassen und Ordnungen des Tierreichs (5)1(7): 1505-1672. Leipzig: Akademische Verlagsgesellschaft Geest & Portig.
Bremer K. 1994. Branch support and tree stability. Cladistics 10: 295-304.
Davie PJF. 2002. Crustacea: Malacostraca Eucarida (Part 2): Decapoda—Anomura, Brachyura. In: Wells A, Houston WWK (eds.). Zoological Catalogue of Australia 19(3B): xiv + 1-641.
Davie PJF, Guinot D. 1996. Two new freshwater crabs in Australocarcinus Davie, with remarks on Trogloplacinae Guinot and Goneplacidae MacLeay (Crustacea: Decapoda: Brachyura). Mem. Qld Mus. 39: 277-287.
D’Udekem d’Acoz C. 1999. Invertaire et distribution des crustacés décapodes de l’Atlantique nord-oriental, de la Méditerranée et des eaux continentals adjacentes au nord de 25 N. Colln Patrim. nat. 40: 1-383.
Felsenstein J. 1985. Confidence limits of phylogenies: an approach using the bootstrap. Evolution 39: 783-791.
Glaessner MF, Secretan S. 1987. Crabes (Crustacea Brachyura) de l’Éocène du Sulaman Range (Pakistan). Ann. Paléont. (Vert.-Invert.) 73: 273-288..
Guinot D. 1969a. Les Goneplacidae: Recherches préliminaires sur les groupements naturels chez les Crustacés, Décapodes, Brachyoures, VII. Bull. Mus. natn Hist. nat. Paris (2)41: 241-265.
Guinot D. 1969b. Les Goneplacidae (suite et fin): Recherches préliminaires sur les groupements naturels chez les Crustacés, Décapodes, Brachyoures, VII. Bull. Mus. natn Hist. nat. Paris (2)41: 688-724.
Guinot D. 1971. Synthèse et bibliographie: Recherches préliminaires sur les groupements naturels chez les Crustacés, Décapodes, Brachyoures, VIII. Bull. Mus. natn Hist. nat. Paris (2)42 (1970): 1063-1090.
Guinot D. 1978. Principes d’une classification évolutive des Crustacés Décapodes Brachyoures. Bull. Biol. Fr. Belg. 112: 211-292.
Guinot D, Macpherson E. 1987. Révision du genre Pilumnoides Lucas, 1844, avec description de quatre espèces nouvelles et création de Pilumnoidinae subfam. nov. (Crustacea Decapoda Brachyura). Bull. Mus. natn Hist. nat. Paris (4)9(1): 211-247.
Karasawa H, Kato H. in press. The family Goneplacidae MacLeay, 1838 (Crustacea: Decapoda: Brachyura): systematics, phylogeny, and fossil records. Paleont. Res. 7.
Lemaitre R, García-Gómez J, von Sternberg R, Campos NH. 2001. A new genus and a new species of crab of family Goneplacidae MacLeay, 1838 (Crustacea: Decapoda: Brachyura) from the tropical western Pacific. Proc. biol. Soc. Wash. 114: 951-963.
Maddison WP, Maddison DL. 2002. MacClade version 4.05. Massachusetts: Sinauer Associates.
Manning RB, Holthuis LB. 1981. West African Brachyuran crabs. Smiths. Contr. Zool. 306: 1-379.
Martin JW, Davis GB. 2001. An updated classification of the recent Crustacea. Nat. Hist. Mus. Los Angeles Cty, Sci. Ser. 39: 1-124.
Ng PKL. 1987. The Indo-Pacific Pilumnidae II. A revision of the genus Rhizopa Stimpson, 1858, and the status of the Rhizopinae Stimpson, 1858 (Crustacea, Decapoda, Brachyura). Indo-Malayan Zool. 4: 69-111.
Ng PKL. 2002. The Indo-Pacific Pilumnidae XVI. On the identity of Pilumnus cristimanus A. Milne Edwards, 1873, and the status of Parapilumnus Kossmann, 1877 (Crustacea: Decapoda: Brachyura), with description of a new species from rubble beds in Guam. Micronesica 34: 209-226.
Ng PKL, Guinot D. 1999. On a new species of deep-water crab of the genus Progeryon (Decapoda, Brachyura, Geryonidae) from Hawai. Crustaceana 72: 685-692.
Ng PKL, Liao LM. 2002. On a new species of Euryozius Miers, 1886 (Crustacea: Decapoda: Brachyura: Pseudoziidae) from the Philippines with notes on the taxonomy of the genus. Proc. biol. Soc. Wash. 115: 585-593.
Rice AL. 1980. Crab zoeal morphology and its bearing on the classification of the Brachyura. Trans. zool. Soc. Lond. 35: 271-424.
Schweitzer CE. 2000. Tertiary Xanthoidea (Crustacea: Decapoda: Brachyura) from the west coast of North America. Jour. Crust. Biol. 20: 715-742.
Schweitzer CE, Feldmann RM, Gonzáles-Barba G, Vega FJ. 2002. New crabs from the Eocene and Oligocene of Baja California Sur, Mexico and an assessment of the evolutionary and paleobiogeographic implications of Mexican fossil decapods. Mem. Paleont. Soc. 59: 1-43.
Serène R, Soh CL. 1976. Brachyura collected during the Thai-Danish Expedition (1966). Phuket mar. Biol. Center, Res. Bull. 12: 1-37.
Serène R. 1964. Redescription du genre Megaesthesius Rathbun et definition des Chasmocarcininae, nouvelle sous-fa-mille des Goneplacidae (Decapoda, Brachyura). Crustaceana 7: 175-187.
Sternberg R von, Cumberlidge N. 2001. Notes on the position of the freshwater crabs within the brachyrhynchan Eubrachyura (Crustacea: Decapoda: Brachyura). Hydrobiologia 449: 21-39.
Swofford DL. 1999. PAUP*: Phylogenetic Analysis Using Parsimony, Version 4b. Illinois: Illinois Natural History Survey.
We thank P.K.L. Ng (National University of Singapore) for valuable comments on some goneplacid taxa and on our phylogenetic analysis, and D. Guinot (Muséum national d’Histoire naturelle, Paris) and Z. Stevcic (Center for Marine Research, Croatia) for their much appreciated comments on some taxa.
Systematic list of genera currently assigned to the Goneplacidae. Asterisks indicate extinct genus (modified from Karasawa & Kato, in press).
Subfamily Mathildellinae Karasawa & Kato, in press (Paleogene-Recent):
Beuroisia Guinot & Richèr de Forges, 1981, Branchioplax Rathbun, 1916*, Intesius Guinot & Richèr de Forges, 1981, Mathildella Guinot & Richèr de Forges, 1981, Neopilumnoplax Serène in Guinot, 1969, Platypilumnus Alcock, 1894, Tehuacana Stenzel, 1944*
Subfamily Euryplacinae Stimpson, 1871 (Eocene-Recent):
Chlinocephalus Ristori, 1886*, Corallicarcinus Müller & Collins, 1991*, Euryplax Stimpson, 1859, Eucrate de Haan, 1835, Fravillea A. Milne Edwards, 1880, Heteroplax Stimpson, 1858, Machaerus Leach, 1818, Nancyplax Lemaitre et al., 2001, Orbitoplax Tucker & Feldmann, 1990*, Psopheticoides Sakai, 1969, Stoaplax Vega et al., 2001*, Trizocarcinus Rathbun, 1914 Viaplax Karasawa & Kato, in press*
Subfamily Goneplacinae MacLeay, 1838 (Late Cretaceous-Recent):
Bathyplax A. Milne Edwards, 1880, Carcinoplax H. Milne Edwards, 1852, Goneplax Leach, 1814, Icriocarcinus Bishop, 1988*, Neommatocarcinus Takeda & Miyake, 1969, Notonyx A. Milne Edwards, 1873, Ommatocarcinus White, 1852, Psopheticus Wood-Mason, 1892, Singhaplax Serène & Soh, 1976
Subfamily Carinocarcinoidinae Karasawa & Kato, in press (Eocene-Oligocene):
Bicarinocarcinus Glaessner & Secretan, 1987*, Carinocarcinoides Karasawa & Fudouji, 2000*
Subfamily Trogloplacinae Guinot, 1986 (Recent):
Australocarcinus Davie, 1987, Trogloplax Guinot, 1986
Subfamily Chasmocarcininae Serène, 1964 (Eocene-Recent):
Camatopsis Alcock & Anderson, 1899, Chasmocarcinus Rathbun, 1898, Chasmocarcinops Alcock, 1900, Collinsius Karasawa, 1993*, Falconoplax van Straelen, 1933*, Gillcarcinus Collins & Morris, 1978*, Hephthopelta Alcock, 1899, Megaesthesius Rathbun, 1909, Mioplax Bittner, 1884*, Orthakrolophos Schweitzer & Feldmann, 2001*, Scalopidia Stimpson, 1858
Goneplacidae incertae sedis (Recent):
Progeryon Bouvier, 1922, Raoulia Ng, 1987, Typhlocarcinodes Alcock, 1900