Translator Disclaimer
30 June 2003 The family Goneplacidae (Crustacea: Decapoda: Brachyura): systematics, phylogeny, and fossil records
Author Affiliations +
Abstract

A phylogenetic analysis of 14 genera of the family Goneplacidae MacLeay (Decapoda: Brachyura: Xanthoidea) is presented based upon 45 adult morphological characters. Two most-parsimonious trees were obtained (length = 87, CI = 0.6667, RI = 0.8242, RC = 0.5495). The present analysis suggests that the Goneplacidae is divided into six subfamilies: Carinocarcinoidinae subfam. nov., Chasmocarcininae Serène, Euryplacinae Stimpson, Goneplacinae MacLeay, Mathildellinae subfam. nov., and Trogloplacinae Guinot. The Carcinoplacinae H. Milne Edwards is synonymised with the Goneplacinae. The family and six subfamilies are defined or redefined based upon the phylogenetic analysis. 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. Our analysis supports recognition of the family Pseudoziidae Alcock by Ng and Liao and suggests that it is the sister to the Eriphiidae MacLeay. A reexamination of fossil records of the Goneplacidae shows that 62 species, 20 genera, and five subfamilies are recognized as fossils. A new monotypic genus Viaplax (Euryplacinae) is erected for Pilumnoplax urpiniana Via. Chlinocephalus Ristori and Gillcarcinus Collins and Morris are moved to the Goneplacidae. Paleopsopheticus Hu and Tao is synonymised with Psopheticus Wood-Mason. Glaessneria Takeda and Miyake is here the junior synonym of Goneplax. Eleven extinct genera previously assigned to the Goneplacidae are not referred to any subfamilies and are transferred out of the Goneplacidae. New combinations include: Carcinoplax proavita (Glaessner), Goneplax arenicola (Glaessner), Euphylax zariquieri (Via) (Portunidae Rafinesque), and Psopheticus shujienae (Hu and Tao).

Introduction

The family Goneplacidae MacLeay, 1838 (Brachyura: Heterotremata: Xanthoidea) has been traditionally recognized as a monophyletic group containing the five subfamilies, Carcinoplacinae H. Milne Edwards, 1852, Eucratopsinae Stimpson, 1871 (= Prionoplacinae Alcock, 1900), Goneplacinae MacLeay, 1838, Hexapodinae Miers, 1886, and Rhizopinae Stimpson, 1858 (Balss, 1957; Sakai, 1976). Guinot (1969a) suggested that the Goneplacidae sensu Balss (1957) was a polyphyletic group and first divided the Goneplacidae into three major groups; “Goneplacidae dérives des Xanthidae”, “Goneplacidae euryplaciens (Euryplacinae)”, and “Goneplacidae carcinoplaciens-gonéplaciens (Carcinoplacinae + Goneplacinae)”. The subfamily Rhizopinae was removed to the Pilumnidae Samouelle, 1819 (Guinot, 1969c, 1978; Ng, 1987; Davie and Guinot, 1996), the Eucratopsinae was assigned to the Panopeidae Ortmann, 1893 (Guinot, 1978; Martin and Abele, 1986) and the Hexapodinae was treated as a family (Guinot, 1978; Manning and Holthuis, 1981). After Balss's (1957) work, two new subfamilies, Chasmocarcininae Serène, 1964 and Trogloplacinae Guinot, 1986, were added to the family. Ng and Wang (1994) moved the Pseudoziinae Alcock, 1898, from the Eriphiidae MacLeay, 1838, to the Goneplacidae. Therefore, the Goneplacidae is now represented by six subfamilies (Lemaitre et al., 2001; Hsueh and Huang, 2002). Subsequently, Ng and Liao (2002) treated the Pseudoziinae as a distinct family.

Glaessner (1969) recognized 20 genera of die Goneplacidae as fossils and assigned 11 extinct genera to the family. Since then, 12 extinct genera have been added:

Caprocancer Müller and Collins, 1991a; Corallicarcinus Müller and Collins, 1991a; Carinocarcinoides Karasawa and Fudouji, 2000; Chumaoia Hu and Tao, 1996; Orthakrolophos Schweitzer and Feldmann, 2001a; Eoplax Müller and Collins, 1991a; Lobogalenopsis Müller and Collins, 1991a; Orbitoplax Tucker and Feldmann, 1990; Paleopsopheticus Hu and Tao, 1996; Paracorallicarcinus Tessier et al., 1999; Pregeryona Hu and Tao, 1996; and Stoaplax Vega et al., 2001. Karasawa and Kato (2001) moved two extinct genera, Maingrapsus Tessier et al., 1999 and Palaeograpsus Bittner, 1875, from the Grapsidae MacLeay, 1838, to the Goneplacidae. They also referred Telphusograpsus Lörenthey, 1902, to the family. Among these, Carinocarcinoides and Stoaplax were referred to the Carcinoplacinae (Karasawa and Fudouji, 2000; Vega et al., 2001), Orbitoplax to the Euryplacinae (Tucker and Feldmann, 1990), and Orthakrolophos to the Chasmocarcininae (Schweitzer and Feldmann, 2001a). Remaining genera were not assigned to any subfamiles within the Goneplacidae because most genera were represented by only carapace specimens. Distinction between the goneplacid genera, and panopeid, pilumnid, and pseudorhombilid genera is difficult based solely upon carapace characters (Schweitzer, 2000).

The first aim of this paper is to provide an adultmorphology-based phylogenetic analysis for 14 genera within the Goneplacidae. A new classification and diagnoses of six subfamilies are presented based upon the phylogenetic analysis. The second aim of this paper is to review fossil taxa previously assigned to the family. All known fossil species and genera within the Goneplacidae are listed.

Phylogenetic analysis of family Goneplacidae

Materials and methods

Fourteen genera including one extinct genus, Carinocarcinoides, within the Goneplacidae, were examined. The analysis also includes Epixanthus Heller, 1861 (Eriphiidae MacLeay, 1838: Oziinae Dana, 1851), Pilumnus Leach, 1815 (Pilumnidae Samouelle, 1819: Pilumninae Samouelle, 1819), and Pseudozius Dana, 1851 (Pseudoziidae Alcock, 1898: Pseudoziinae Alcock, 1898) as ingroup taxa to analyze a sister-group relationship of the Goneplacidae. The analyses were based upon the examination of material deposited in the Kanagawa Prefectural Museum of Natural History, Odawara, Japan; the Mizunami Fossil Museum, Mizunami, Japan; the Natural History Museum and Institute, Chiba, Japan; and the National Museum of Natural History, Smithsonian Institution, Washington D.C., U.S.A. The material examined is listed in Table 1. If material was unavailable, the descriptive information of taxa was obtained from the literature. The subfamilial arrangement of the genera conforms to Guinot (1970 [1971]), Guinot and Richer de Forges (1981), Serène (1984), Davie and Guinot (1996), Karasawa and Fudouji (2000), and Ng and Liao (2002).

Table 1

Taxa included in the analysis. Abbreviations: CBM, Natural History Museum and Institute, Chiba; KPM, Kanagawa Prefectural Museum of Natural History; MFM, Mizunami Fossil Museum; NMNH, National Museum of Natural History, Smithsonian Institution; *1, Guinot (1989); *2, Guinot (1990); *3, Ikeda (1998), *4, Rathbun (1918); *5, Felder and Rabalais (1986); *6, Guinot (1986); *7, Guinot and Richer de Forges (1981).

i1342-8144-7-2-129-t01.gif

An outgroup was chosen to polarize the character states. The Goneplacidae does not have a reliable sister group. Ortmann (1893) thought that the Goneplacidae (= his Carcinoplacidae + Goneplacidae) were derived from the Eriphiidae (= his Menippidae). Guinot (1969c) and Stevcic in Martin and Davis (2001) mentioned that there is a close relationship between the Goneplacidae and Geryonidae Colosi, 1923 based upon adult morphology. Rice (1980) and Martin (1984) showed that the family is most similar to the Pilumnidae based upon zoeal morphology. Von Sternberg and Cumberlidge (2001) suggested based upon cladistic and phenetic analysis that the Goneplacidae may be more closely related to the Portunidae Rafinesque, 1815, than to any families of the Xanthoidea. Therefore, the cladogram was rooted against a “hypothetical ancestor”. Table 2 lists 45 adult morphological characters and character states used in the analysis. The missing data were scored as unknown. The data matrix is provided in Table 3. Forty-five characters were included in the data matrix (Table 3). There are 42 binary characters and three multistate characters. In the text, characters and character states are indicated by numbers in parentheses (e.g., 1-0 = character 1 + character state 0).

Table 2

Characters and their states used in the phylogenetic analysis.

i1342-8144-7-2-129-t02.gif

Table 3

Input data matrix of 45 characters and 17 genera. Missing character states are shown by?.

i1342-8144-7-2-129-t03.gif

The phylogenetic analysis used PAUP* 4.0b (Swofford, 1999), utilizing a data matrix originating in MacClade version 4.03 (Maddison and Maddison, 2001). Heuristic search analyses were performed with the following options in effect: addition sequence, 100 replications with random input order; one tree held at each step during stepwise addition; tree-bisection-reconnection (TBR) branch stepping performed; MulTrees option activated; steepest descent option not in effect; branches having maximum length zero collapsed to yield polytomies; topological constraints not enforced; tree unrooted; multistate taxa interpreted as polymorphism; character state optimization; and accelerated transformation (ACCTRAN). All characters were unordered, unsealed and equally weighted. Relative stability of clades was assessed using bootstrap (Felsenstein, 1985) and decay analyses (Bremer, 1994). The bootstrapping was based on 100 replicates of random input order. The Bremer support was obtained using constraint trees generated by AutoDecay 4.02 (Eriksson, 1999) and analyzed using PAUP*.

Results

The present analysis yielded two most-parsimonious trees, 87 steps long with a consistency index (CI) of 0.6667, a retention index (RI) of 0.8242 and a rescaled consistency index (RC) of 0.5495. A strict consensus tree of two most-parsimonious trees, indicating bootstrap and Bremer support, is given in Figure 1. Fourteen distinct clades are recognized. Each clade is numbered with character state changes in Figure 2.

Figure 1

Strict consensus tree of two most-parsimonious trees of 14 genera within the Goneplacidae. Length = 87, Consistency index = 0.6667, Retention index = 0.8242, Rescaled consistency index = 0.5495. Numbers above branches are Bootstrap support and numbers below branches are Bremer support.

i1342-8144-7-2-129-f01.gif

Figure 2

Strict consensus tree of two most-parsimonious trees of 14 genera within the Goneplacidae. Length = 87, Consistency index = 0.6667, Retention index = 0.8242, Rescaled consistency index = 0.5495. Character changes are indicated. Numbers above branches are clade numbers.

i1342-8144-7-2-129-f02.gif

Clade 1:

Epixanthus + Pseudozius (Eriphiidae + Pseudoziidae). In the examined material Pseudozius and Epixanthus are sister taxa nested as the most basal clade. This clade, with 65% bootstrap support and Bremer support of 2, is united by three synapomorphies (6-1, 7-1, 32-1). None is unique.

Clade 2:

Pilumnus + Goneplacidae. Pilumnus and taxa of the Goneplacidae clade, with 69% bootstrap support and Bremer index of 2, share four synapomorphies, two of which are unique and never reversed: the absence of frontal teeth (3-1) and the absence of a median sulcus on the thoracic sternite 4 (24-1).

Clade 3:

Goneplacidae. The monophyly of the Goneplacidae, with 77% bootstrap support and Bremer index of 3, is well defined by seven synapomorphies, three of which are unique and unreversed: a wide thoracic sternum (21-1), a medially interrupted sulcus delimiting thoracic sternites 6 and 7 (22-1), and long meri of pereiopods 2–5 (42-1).

Clade 4:

Mathildella + Beuroisia + Intesius (Goneplacidae incertae sedis). The Mathildella + Beuroisia + Intesius clade, with 93% bootstrap support and Bremer support of 3, is unambiguously united by four synapomorphies: laterally broad antennular fossae (10-1), the absence of a prolongation of the thoracic sternite 7 in male (26-0; reversal), a strongly inflated basal part of male gonopod 1 (35-1), and a spatulate dactylus of pereiopods 5 (44-1). Two synapomorphies (10-1, 35-1) are unique and never reversed.

Clade 5:

Mathildella + Beuroisia. Only one unique synapomorphy, a semicircular male telson (15-1), defines this clade.

Clade 6:

Euryplacinae + Carcinoplacinae + Goneplacinae + Carinocarcinoides + Trogloplacinae + Chasmocarcininae. This clade, with 85% bootstrap support and Bremer support of 2, shares eight synapomorphies, four of which are unique and never reversed: a medially interrupted sulcus delimiting thoracic sternites 7 and 8 (23-1), an anterior margin of the male sterno-abdominal cavity reaching the anterior part of the thoracic sternite 4 (25-1), the absence of dark-colored cheliped fingers (40-1) and the possession of dactyli of pereiopods 2–5 which terminate with acute chitinous tips (43-1).

Clade 7:

Eucrate + Euryplax + Heteroplax (Euryplacinae). The Euryplacinae clade has 76% bootstrap support and Bremer support of 2. Seven synapomorphies (5-2, 11-0, 14-1, 16-1, 31-1, 36-1, 37-1) well define this clade. A unique synapomorphy is distinctly narrow male abdominal somites 4-6 (16-1). One synapomorphy, the presence of the basal article of antenna reaching the front (11-0), is a reversal.

Clade 8:

Euryplax + Heteroplax. Only one synapomorphy, a long eye stalk (9-1), defines this clade.

Clade 9:

Carcinoplacinae + Goneplacinae + Carinocarcinoides + Trogloplacinae + Chasmocarcininae. This clade, with 80% bootstrap support and Bremer index of 2, shares four synapomorphies: the supraorbital angle fused to the frontal margin (4-1), the absence of upper orbital fissures (6-1), a sinuous gonopod 1 (32-0; reversal), and the absence of marginal setae of dactyli of pereiopods 5 (45-0; reversal). The supraorbital angle fused to the frontal margin (4-1) is a unique synapomorphy.

Clade 10:

Psopheticus + Carcinoplax + Ommatocarcinus + Goneplax (Carcinoplacinae + Goneplacinae). The Carcinoplacinae + Goneplacinae clade, with 72% bootstrap support and Bremer index of 2, is well defined by three synapomorphies, two of which are unique: the absence of a median notch on the frontal margin (1-1), and a truncated apex of gonopod 1 (34-1) and the possession of a ventral spine of the cheliped carpus (41-1). The sister-group relationship of the clade (Psopheticus, Carcinoplax and Ommatocarcinus + Goneplax) remained unresolved.

Clade 11:

Ommatocarcinus + Goneplax (Goneplacinae). Five synapomorphies support this clade. Only one synapomorphy, the possession of the front with a median projection (2-1), is unique. Two synapomorphies, the male abdominal somite 2, which is much narrower than somite 3 (18-0), and the absence of a ventral spine of the carpus of the cheliped (41-0), are reversals.

Clade 12:

Carinocarcinoides + Trogloplacinae + Chasmocarcininae. This clade, with 91% bootstrap support and Bremer index of 3, shares three unique synapomorphies: the possession of fused male abdominal somites 3–5 (20-1), the thoracic sternite 8 overlying posterolaterally sternite 7 (27-1), and the thoracic sternite 8 which is visible ventrally (29-1).

Clade 13:

Trogloplacinae + Chasmocarcininae. The Trogloplacinae + Chasmocarcininae clade, with 88% boot-strap support and Bremer support of 5, is evidently united by five synapomorphies (5-0, 8-2, 12-1, 17-1, 18-0, 28-1). Three of these synapomorphies, the presence of maxilliped 3 ischium about equal to merus (12-1), male abdominal somite 3 much narrower than thoracic sternites 7 and 8 (17-1), and the presence of a supplementary plate of male thoracic sternite 8 (27-1), are unique.

Clade 14:

Camatopsis + Chasmocarcinus (Chasmocarcininae). This clade, with 96% bootstrap support and Bremer support of 3, is well defined by four synapomorphies (13-1, 37-1, 44-2, 45-0). The possession of a suboval merus of maxilliped 3 (13-1) is a unique synapomorphy.

Discussion

Guinot (1969a, b, c; 1970 [1971]) divided the family Goneplacidae sensu Balss (1957) into three major groups; “Goneplacidae dérives des Xanthidae”, “Goneplacidae euryplaciens (Euryplacinae)”, and “Goneplacidae carcinoplaciens-gonéplaciens (Carcinoplacinae + Goneplacinae)”. Glaessner (1969) and Sakai (1976) used the classification of the Goneplacidae sensu Balss, while Serène and Soh (1976), Manning and Holthuis (1981), and Williams (1984) partly accepted Guinot's concept for the classification of the family.

Since then, genera belonging to her “Goneplacidae dérives des Xanthidae” were removed to other families. Guinot (1978) and Martin and Abele (1986) transferred the Eucratopsinae to the family Panopeidae. The Rhizopinae sensu lato is currently placed in the Pilumnidae (Guinot, 1969c, 1978; Ng, 1987; Davie and Guinot, 1996). Litocheira Kinaham, 1856 sensu stricto (see Guinot, 1970 [1971]; Türkay, 1975), is referred to her “Goneplacidae pilumniens sensu stricto”, while the genus has not been assigned to any of the pilumnid subfamilies. Guinot (1969c, 1970 [1971]) referred Galene De Haan, 1833, to her “Goneplacidae pilumniens sensu lato”, while Takeda (1976) included the genus within the subfamily Galeninae Alcock, 1898, of the Xanthidae sensu lato. Ng (1998) and Schweitzer (2000) classified Galene within the Pilumnidae, following Guinot (1969c, 1970 [1971]); therefore, species of the Galeninae are thought to be members of the Pilumnidae (Ng et al., 2001; Hsueh and Huang, 2002). The Pseudorhombilinae Alcock, 1900, previously referred to the Goneplacidae, was also included in her “Goneplacidae dérives des Xanthidae” but Hendrickx (1998) treated it as a distinct family.

Davie and Guinot (1996) indicated that the Goneplacidae contains five subfamilies, Goneplacinae MacLeay, Carcinoplacinae H. Milne Edwards, Chasmocarcininae Serène, Trogloplacinae Guinot and Euryplacinae Stimpson. Ng and Wang (1994) transferred the Pseudoziinae Alcock from the Eriphiidae to the Goneplacidae. Therefore, Lemaitre et al., (2001) and Hsueh and Huang (2002) currently divided the Goneplacidae into six subfamilies. Subsequently, Ng and Liao (2002) excluded the Pseudoziinae from the Goneplacidae and treated it as a distinct family.

The present phylogenetic analysis well supports the monophyly of the Goneplacidae as envisioned by Davie and Guinot (1996). Six synapomorphies, three of which are unique and unreversed, well define the Goneplacidae (Figure 2, Clade 3). The present analysis suggests that the Intesius + Mathildella + Beuroisia (Goneplacidae incertae sedis) clade within the Goneplacidae is the most basal, followed by the Euryplacinae, the Carcinoplacinae + Goneplacinae, and the most advanced clade, Carinocarcinoides + Trogloplacinae + Chasmocarcininae.

Pseudozius, the type genus of the Pseudoziidae, is the sister to Epixanthus (Eriphiidae; Oziinae) (Figure 2; clade 1) and both genera are united by three synapomorphies. Alcock (1898) originally placed Pseudozius within his alliance Pseudozioida Alcock (= Pseudoziinae Alcock; nom. transl. of Takeda (1976)) within his Menippinae of the family Xanthidae sensu lato and subsequent workers (i.e., Guinot, 1970[1971]; Sakai, 1976; Takeda, 1976) also placed it within the Xanthidae sensu lato. Crosnier in Serène (1984) referred Pseudozius to incertae sedis within the Menippidae (= Eriphiidae). Ng and Wang (1995) moved the subfamily from the Eriphiidae to the Goneplacidae. Subsequently, Ng and Liao (2002) recognized the Pseudoziinae as a separate family and divided it into two subfamilies, Pseudoziinae and Planopilumninae Serène, 1984. In their work the Pseudoziinae contains four genera, Euryozius Miers, 1886, Flindersoplax Davie, 1989, Platychelonion Crosnier and Guinot, 1969, and Pseudozius, and the Planopilumninae is a monotypic subfamily. Our analysis supports the recognition of the Pseudoziidae by Ng and Liao (2002) and suggests that the family is the sister taxon of the Eriphiidae. Members of the subfamilies Eriphiinae, Oziinae, Menippinae Ortmann, 1893, and Dacryopilumninae Serène, 1984, within the Eriphiidae have a long gonopod 2 with a filamentous, long flagellum (36-0, 37-0, 38-1) while Pseudozius is characterized by having a short gonopod 2 and by lacking a filamentous, long flagellum of gonopod 2 (36-1, 37-1, 38-0).

The most basal Intesius + Mathildella + Beuroisia clade shares four synapomorphies, two of which are unique and never reversed: laterally broad antennular fossae (10-1) and a strongly inflated basal part of gonopod 1 (35-1) (Figure 2; clade 4). The subfamilial placement of three genera has not been well documented. Guinot and Richer de Forges (1981) erected two new genera, Mathildella and Beuroisia, based upon examination of new material and species previously assigned to Neopilumnoplax Serène in Guinot, 1969c, but did not designate subfamilial placement for Mathildella and Beuroisia or for another new genus, Intesius. Guinot (1970 [1971]) placed Neopilumnoplax within “Autres Carcinoplacinae-Goneplacinae” of the Goneplacidae, whereas Sakai (1976) placed it within the Carcinoplacinae. Poupin (1996) assigned Intesius to the Goneplacidae, and Beuroisia and Mathildella to “Xanthoidea incertae sedis”. Ng et al. (2001) and Hsueh and Huang (2002) placed Mathildella within the Carcinoplacinae. The present analysis supports that these three genera should be included within the Goneplacidae. The three genera within this clade differ significantly from other goneplacid genera (Figure 2; clade 6) because they lack the diagnostic synapomorphies of clade 6; therefore, they cannot be placed within previously known subfamilies. A new subfamily, Mathildellinae, is erected herein for these genera.

Several workers did not recognize the Euryplacinae as a valid taxon. Indeed, Balss (1957) included Eucrate and Heteroplax within the Carcinoplacinae, and Euryplax within the Prionoplacinae, and Sakai (1976) classified Eucrate and Heteroplax within the Carcinoplacinae. However, the Euryplacinae (Figure 2, Clade 7) is well supported as monophyletic by seven synapomorphies, one of which is unique and never reversed, distinctly narrow male abdominal somites 4-6 (16-1). The present analysis strongly supports recognition of the subfamily by Guinot (1969a, b, c, 1970 [1971]), Manning and Holthuis (1981), Ng et al. (2001), and Hsueh and Huang (2002). The Euryplacinae clade is the sister to the Carcinoplacinae + Goneplacinae + Carinocarcinoides + Trogloplacinae + Chasmocarcininae clade (Figure 2, Clade 9).

For the Carcinoplacinae, represented by Psopheticus and Carcinoplax, the analysis is unable to resolve the relationships between both taxa and other goneplacines, since they nest in a polytomy with the Goneplacinae clade (Figure 2, Clade 10). In one of the two most-parsimonious trees the subfamily is monophyletic whereas in another tree it is paraphyletic. The monophyly of the Goneplacinae is supported by five synapomorphies, but it is nested among the goneplacine genera (Figure 2, Clade 11). The Carcinoplacinae should either be synonymised with the Goneplacinae or divided into three subfamiles. In the latter scheme, a new monotypic subfamily would have to be proposed for Psopheticus. The Goneplacinae clade with Bremer support of 1 is more weakly defined than the Carcinoplacinae + Goneplacinae clade with Bremer support of 2. In the present analysis the Carcinoplacinae + Goneplacinae clade shares three synapomorphies, two of which are unique: the absence of a median notch on the frontal margin (1-1), and a truncated apex of gonopod 1 (34-1). Therefore, rather than proposing three subfamilies, each with weakly defined synapomorphies and with weak Bremer support, it is considered best to place Carcinoplax and Psopheticus, previously assigned to the Carcinoplacinae, within the Goneplacinae. Our phylogenetic analysis supports Guinot's concept of a “lignée Carcinoplacienne-Gonéplacienne” and “groupement Carcinoplax - Psopheticus - Goneplax - Ommatocarcinus” (Guinot, 1969b, c).

The Carinocarcinoides + Trogloplacinae + Chasmocarcininae clade is characterized by three unique synapomorphies: fused male abdominal somites 3–5 (20-1), thoracic sternite 8 overlying posterolaterally sternite 7 (27-1), and thoracic sternite 8 visible ventrally (29-1) (Figure 2, Clade 12). Karasawa and Fudouji (2000) originally placed Carinocarcinoides within the Carcinoplacinae; however, the present analysis suggests that the genus does not belong to the Carcinoplacinae. Carinocarcinoides is the first to diverge within the clade, characterized by having more or less defined dorsal regions of the carapace (7-0) and elongate chelipeds (39-1). The Trogloplacinae + Chasmocarcininae clade is unambiguously united by six synapomorphies, three of which are unique: ischium of maxilliped 3 about equal to merus (12-1), male abdominal somite 3 much narrower than thoracic sternites 7 and 8 (17-1), and the possession of the supplementary plate of thoracic sternite 8 in males (27-1) (Figure 2, Clade 13). On the basis of the phylogenetic analysis, Carinocarcinoides cannot be included in either subfamily, since it lacks their diagnostic synapomorphies, and the genus is here recognized as the type of a new monotypic subfamily, Carinocarcinoidinae.

The Trogloplacinae is here derived as the sister group to the Chasmocarcininae (Figure 2, Clade 14). Davie and Guinot (1996) suggested that the Trogloplacinae had close affinities with the Chasmocarcininae. We concur. The Trogloplacinae is a weakly defined subfamily lacking the diagnostic synapomorphies of the Chasmocarcininae. The Chasmocarcininae is a distinctive subfamily clearly defined by four autapomorphies (13-1, 37-1, 44-1, 45-1), one of which is unique: the possession of a suboval merus of maxilliped 3 (13-1).

Systematics

Family Goneplacidae MacLeay, 1838 emend.

Diagnosis.

Carapace transversely rectangular, trapezoidal or rounded; dorsal regions weakly distinct or indistinct; front straight, sometimes bearing median notch or median projection, without teeth; notch between frontal and supraorbital angle present or absent; upper orbital margin with or without fissures; anterolateral margin usually toothed; inner antennular septum a thin plate; buccal frame quadrangular; epistome well defined; palp of maxilliped 3 articulating on or near anteromesial corner of merus; exopod wide; male abdomen with all free somites or fused somites 3–5; thoracic sternum wide with all sutures interrupted, rarely with continuous suture delimiting sternites 7 and 8; sternite 4 lacking median sulcus; sternite 7 usually with posterolateral prolongation; chelipeds heterochelate; pereiopods 2–5 long; dactyli of pereiopods 2–5 with or without corneous tips; male genital openings coxal; gonopod 1 stout, sinuous or curved, usually with simple apex; gonopod 2 long or short.

Type genus.

Goneplax Leach, 1814.

Subfamilies included.

Carinocarcinoidinae subfam. nov.; Chasmocarcininae Serène, 1964; Euryplacinae Stimpson, 1871; Goneplacinae MacLeay, 1838; Mathildellinae subfam. nov.; Trogloplacinae Guinot, 1986.

Remarks.

The diagnosis is based upon Balss (1957) and the present phylogenetic analysis.

Subfamily Mathildellinae subfam. nov.

Diagnosis.

Carapace usually flattened with weakly defined dorsal regions; front straight with shallow median notch; supraorbital angle separated from frontal margin; orbit relatively small with upper orbital fissures; anterolateral margin bearing five teeth; eye stalk short; antennular fossae broad laterally; merus of maxilliped 3 subquadrate, much longer than ischium; male abdomen filling entire space between coxae of pereiopods 5, usually with all free somites; thoracic sternum wide with interrupted sutures excluding continuous suture delimiting sternites 7 and 8; sternite 7 without posterolateral prolongation; sterno-abdominal cavity reaching posterior of sternite 4; chelipeds with dark-colored fingers; dactyli of pereiopods 2–5 with corneous tips; dactyli of pereiopods 5 spatulate with setae; gonopod 1 stout, curved, strongly inflated basally, with simple apex; gonopod 2 usually long with long flagellum.

Type genus.

Mathildella Guinot and Richer de Forges, 1981.

Genera included.

Beuroisia Guinot and Richer de Forges, 1981; Branchioplax Rathbun, 1916; Intesius Guinot and Richer de Forges, 1981; Mathildella; Neopilumnoplax Serène in Guinot, 1969; Platypilumnus Alcock, 1894; Tehuacana Stenzel, 1944.

Discussion.

The Mathildellinae is the most basal group within the Goneplacidae based upon the present phylogenetic analysis. The subfamily is well defined by the presence of more or less defined anterior dorsal regions, laterally broad antennular fossae, a complete sulcus delimiting thoracic sternites 7 and 8, an anterior end of the sterno-abdominal cavity located on the posterior half of the thoracic sternite 4, the absence of a posterolateral prolongation of the thoracic sternite 7, the presence of dark-colored cheliped fingers, dactyli of pereiopods 2–5 with corneous tips, and a strongly inflated basal part of gonopod 1, all of which other goneplacid subfamilies lack.

Alcock (1900) questionably referred Platypilumnus to the Goneplacidae. Guinot (1970 [1971]) placed Platypilumnus within the Geryonidae, while Manning and Holthuis (1989) did not include the genus within the Geryonidae. Richer de Forges (1996) showed that Platypilumnus has close affinities with Neopilumnoplax and Intesius. We concur. Crosnier and Guinot (1969) suggested that Platychelonion is similar to Neopilumnoplax whereas Guinot (1970 [1971]) questionably referred it to the Geryonidae. Manning and Holthuis (1989) excluded the genus from the Geryonidae and Davie (1989) suggested that the genus bears a close resemblance to the Goneplacidae. We place Platychelonion within the Pseudoziidae, following Ng and Liao (2002).

Tucker and Feldmann (1990), Schweitzer et al. (2000), and Schweitzer (2000) described well preserved specimens of Branchioplax washingtoniana Rathbun, 1916, the type species of Branchioplax, from Palaeogene rocks of the U.S.A. In her taxonomic review of Branchioplax Schweitzer (2000) synonymised Pilumnoplax hannibalanus Rathbun, 1926, with B. washingtoniana and moved Pilumnoplax carmanahensis Rathbun, 1926, to Branchioplax. Balss (1957) and Glaessner (1969) referred Branchioplax to the Carcinoplacinae, while Schweitzer (2000) suggested that the genus is similar to the extant Chacellus Guinot, 1969c. Chacellus is now placed within the family Pseudorhombilidae (Hendrickx, 1998). Examination of illustrations of B. washingtoniana and B. pentagonalis (Yokoyama, 1911) indicates that the genus belongs to the Mathildellinae. In Branchioplax the anterior dorsal regions are more or less defined; a nearly straight frontal margin bears a median notch; the supraorbital angle is developed; the upper orbital margin possesses two fissures; the anterior end of the sterno-abdominal cavity located on the posterior part of sternite 4; the sulcus delimiting thoracic sternites 7 and 8 is complete; the posterolaterally directed prolongation of thoracic sternite 7 is not developed; and the male abdomen consists of seven free somites. These characters are also definitive characters of the sub-family.

Stenzel (1944) established the monotypic genus Tehuacana based upon a male specimen from the Palaeogene of the U.S.A. and compared this new genus with “Pilumnoplax Stimpson, 1858”. This genus has upper orbital fissures and more or less defined anterior dorsal regions, and lacks a prolongation of thoracic sternite 7. Therefore, the genus is here assigned to the Mathildellinae.

Fossil records.

Fossil records of the Mathildellinae are represented by two extinct genera known from the Palaeogene (Table 4).

Table 4

Distributions and geologic ranges of recognized fossil species of the subfamily Mathildellinae.

i1342-8144-7-2-129-t04.gif

Subfamily Euryplacinae Stimpson, 1871 emend.

Diagnosis.

Carapace usually with poorly defined dorsal regions; front straight with shallow median notch; supraorbital angle distinct; orbit sometimes deep, large, with upper orbital fissures; anterolateral margin bearing two to five spines; eye stalk short or long; basal article of antenna reaching front; merus of maxilliped 3 subquadrate, much longer than ischium; male abdomen filling entire space between coxae of pereiopods 5, with all free somites; somites 4–6 much narrower than somite 3; telson usually longer than wide; thoracic sternum wide with sutures all interrupted; sternite 7 with posterolateral prolongation; sternite 8 visible in posterior view; sterno-abdominal cavity reaching anterior of sternite 4; chelipeds without darkcolored fingers; dactyli of pereiopods 2–5 without corneous tips; dactyli of pereiopods 5 usually styliform, with or without setae; gonopod 1 stout, curved, with simple apex; gonopod 2 very short with short flagellum.

Type genus.

Euryplax Stimpson, 1859.

Genera included.

Chlinocephalus Ristori, 1886; Corallicarcinus Müller and Collins, 1991a; Euryplax; Eucrate De Haan, 1835; Fravillea A. Milne Edwards, 1880; Heteroplax Stimpson, 1858; Machaerus Leach, 1818; Nancyplax Lemaitre et al., 2001; Orbitoplax Tucker and Feldmann, 1990; Psopheticoides Sasaki, 1969; Stoaplax Vega et al., 2001; Trizocarcinus Rathbun, 1914; Viaplax gen. nov.

Discussion.

Balss (1957) and Sakai (1976) placed members of the subfamily within the Carcinoplacinae, while Guinot (1970 [1971]), Serène and Soh (1976), Manning and Holthuis (1981), Williams (1984), Ng et al. (2001), and Hsueh and Huang (2002) indicated that the Euryplacinae is a valid taxon. The present analysis strongly supports the monophyly of the Euryplacinae.

Ristori (1886) described a new genus and species, Chlinocephalus demissifrons, from the Pliocene of Italy and originally placed it within the Cancridae Latreille, 1802. Glaessner (1929) referred Chlinocephalus to the Goneplacidae and in 1969 removed the genus to the Xanthidae sensu lato. The genus is reassigned to the Euryplacinae because the male abdomen consists of seven free somites, the telson of the male abdomen is much longer than wide, and the abdominal somites 4 and 5 are much narrower than the somite 3. The genus may resemble Eucrate, but differs by the presence of transverse ridges of the dorsal surface.

Via (1959) described a new species, Pilumnoplax urpiniana from the Eocene of Spain. Feldmann and Maxwell (1990) referred this species to Carcinoplax and Schweitzer (2000) assigned it to the Pilumnidae. Via (1959, 1969) indicated that Pilumnoplax urpiniana has three anterolateral teeth, while in the species a broken fourth anterolateral tooth (Via, 1969, pl. 36, figs. 2, 2b) is observed. Pilumnoplax urpiniana possesses carapace and male abdomen characters most like those of members of Eucrate; the front has a median notch; the supraorbital angle is well marked; the upper orbital margin bears two shallow notches; the anterolateral margin has four anterolateral teeth; the male abdominal somites 5 and 6 are much narrower than somite 3; and the telson is much longer than wide. However, the species differs from species of Eucrate by having a flattened dorsal surface, well developed triangular anterolateral teeth, more or less defined cervical and branchiocardiac grooves, and well marked epibranchial regions. Therefore, Viaplax gen. nov. is here erected with a type species, Pilumnoplax urpiniana Via, 1959. The generic name is derived from the late Dr. L. Via, a Spanish paleontologist, and the suffix-plax (flat), which is used in names of related genera. The gender is feminine. The carapace of the new genus may also be similar to that of Benthopanope Davie, 1989, of the pilumnid Heteropanopinae Alcock, 1898, nom. transl. herein pro Heteropanopeinae Alcock, 1898, nom. transl. Serène (1984)). It is readily distinguished from Benthopanope by the possession of a wider orbital margin and well developed anterolateral teeth, and the absence of granular dorsal crests on the carapace and prominent median lobes on the frontal margin.

Vega et al. (2001) erected a new monotypic genus, Stoaplax, containing S. nandachare Vega et al., 2001, from the middle Eocene of México. They referred the genus to the Carcinoplacinae and indicated that it is most similar to Orbitoplax. Herein, the genus is removed to the Euryplacinae by having a medially notched frontal margin and a wide upper orbital margin with a fissure.

The Eocene genus Corallicarcinus possesses carapace characters like those of the extant Euryplax; however, in Corallicarcinus the carapace has distinct ridges extending onto the dorsal surface from the second and third anterolateral teeth.

Fossil records.

Seven genera including five extinct ones are known from the fossil record (Table 5). Four extinct genera are known from the Eocene and one is from the Pliocene. The geologic range of two extant genera, Eucrate and Euryplax, extend back to the Oligocene.

Table 5

Distributions and geologic ranges of recognized fossil species of the subfamily Euryplacinae. Asterisk indicates extant species.

i1342-8144-7-2-129-t05.gif

Subfamily Goneplacinae MacLeay, 1838 emend.

(= Subfamily Carcinoplacinae H. Milne Edwards, 1852)

Diagnosis.

Carapace with poorly defined dorsal regions; front straight without median notch, sometimes with low median projection; notch between frontal margin and supraorbital angle indistinct; orbit without upper orbital fissures; anterolateral margin bearing one to three spines; eye stalk short or long; merus of maxilliped 3 subquadrate, much longer than ischium; male abdomen filling entire space between coxae of pereiopods 5, with all free somites; thoracic sternum wide with sutures all interrupted; sternite 7 with posterolateral prolongation; sternite 8 visible in posterior view; sterno-abdominal cavity reaching anterior of sternite 4; chelipeds usually with lateral spine on carpus and without dark-colored fingers; dactyli of pereiopods 2–5 without corneous tips; dactyli of pereiopod 5 styliform or spatulate, with or without setae; gonopod 1 stout, sinuous, usually with truncated apex; gonopod 2 usually long with long flagellum.

Type genus.

Goneplax Leach, 1814.

Genera included.

Bathyplax A. Milne Edwards, 1880; Carcinoplax H. Milne Edwards, 1852; Goneplax; Neom-matocarcinus Takeda and Miyake, 1969; Ommatocarcinus White, 1852; Psopheticus Wood-Mason, 1892; Singhaplax Serène and Soh, 1976.

Discussion.

The Goneplacinae was previously distinguished from the Carcinoplacinae by the following characters: carapace subquadrate in outline, greatest carapace width is at the outerorbital angle, front is usually narrow, and orbit is extremely elongate (Balss, 1957; Sakai, 1976; Hsueh and Huang, 2002). However, the Carcinoplacinae is herein recognized as a synonym of the Goneplacinae based upon the present analysis.

Via (1959) described a new species, Ommatocarcinus zariquieri, from the Eocene of Italy. In his 1969 work the present species was well figured. This species is not a member of Ommatocarcinus because the dorsal carapace possesses three well defined transverse ridges (Via, 1969, pl. 37, figs. 1, 1a, 2, 2a), the maximum carapace width is at the anterolateral angle.(Via, 1969, pl. 37, figs. 1, 1a, 2, 2a), there is a well defined median groove on thoracic sternites 2 and 3 (Via, 1969, pl. 37, fig. 1b), and male abdominal somites 3–5 are fused (Via, 1969, pl. 37, fig. 1b). In the possession of fused abdominal somites 3–5 Takeda and Miyake (1969) and Jenkins (1975) noted that the species may be transferred to Neommatocarcinus Takeda and Miyake, 1969, but O. zariquieri apparently differs from the sole included species of Neommatocarcinus, N. huttoni (Filhol, 1885), by carapace and thoracic sternum characters. The species is here assigned to Euphylax Stimpson, 1860, of the family Portunidae Rafinesque, based upon the characters discussed above. In Ommatocarcinus there is only one transverse ridge on the dorsal surface, the maximum carapace width is at the outerorbital angle, a median groove is absent on thoracic sternites 2 and 3, and all male abdominal somites are free.

Glaessner (1960) described Ommatocarcinus arenicola Glaessner, 1960, from the lower Miocene of New Zealand and noted that “The new species is closer in the shape of its carapace to the less specialized genus Goneplax Leach, but the front is more like that of Ommatocarcinus, to which the species is assigned as an early primitive form”. Takeda and Miyake (1969) proposed a new goneplacid genus, Glaessneria, for the species. We cannot concur. The carapace in this species bears two anterolateral teeth with an anterolaterally directed outerorbital tooth, has a relatively wide front without a median projection, and lacks a distinct transverse ridge dorsally; therefore, the species is here moved to Goneplax. Glaessneria thus becomes a junior subjective synonym of Goneplax.

Paleopsopheticus Hu and Tao, 1996, is a junior subjective synonym of Psopheticus. Hu and Tao (1996) distinguished the present monotypic genus from Psopheticus by having a small-sized carapace, a nearly straight anterolateral margin, a rounded posterior margin, and equal-sized anterolateral spines (modified from Hu and Tao, 1996, p. 102). We believe that these characters cannot define the genus but define Psopheticus shujenae (Hu and Tao, 1996) comb. nov. Examination of their figures (Hu and Tao, 1996, pl. 49, figs. 1, 2, 5, 6) suggests that the species is quite similar to most members of Psopheticus (i.e. P. stridulans Wood-Mason, 1892, P. vocans Guinot, 1985).

Glaessner (1960) described a new species, Galene proavita, from the Miocene of New Zealand. This species is here moved to Carcinoplax because the carapace has smooth, poorly defined anterior dorsal regions with a straight frontal margin, the thoracic sternum and male abdomen are wide, and the telson of the male abdomen is about as long as wide. In members of Galene the frontal margin is bilobed; the anterior mesogastric process is more or less defined; the cardiac region is longer than wide; the thoracic sternum is much longer than wide with a narrow sterno-abdominal cavity; the male abdominal somites 4–6 are much narrower than somite 3 with a long, elongate telson. Pilumnoplax petrificus Hu and Tao, 1996, from the Pleistocene of Taiwan, is identical with Carcinoplax prisca Imaizumi, 1961, because the carapace is rounded-hexagonal in outline and slightly wider than long, and a large outerorbital tooth is directed sharply forwards. Hu and Tao (1996) described a new species, Carcinoplax linae, from the upper Pliocene of Taiwan. They compared the species with Carcinoplax longimanus (De Haan, 1833) rather than with the quite similar extant species Carcinoplax purpurea Rathbun, 1914, and in fact no substantive difference between C. linae and C. purpurea can be found. Carcinoplax linae is here regarded as a junior synonym of C. purpurea.

Fossil records.

Four genera, Carcinoplax, Goneplax, Ommatocarcinus, and Psopheticus, are recognized as fossils (Table 6). Most species of these genera are known from the Neogene and two species, Carcinoplax temikoensis and Psopheticus shujense, are from the Palaeogene.

Table 6

Distributions and geologic ranges of recognized fossil species of the subfamily Goneplacinae. Asterisk indicates extant species.

i1342-8144-7-2-129-t06.gif

Subfamily Carinocarcinoidinae subfam. nov.

Diagnosis.

Carapace with more or less defined dorsal regions; front straight without median notch; notch between frontal margin and supraorbital angle indistinct; upper orbital margin without fissures; anterolateral margin bearing three spines; merus of maxilliped 3 subquadrate, much longer than ischium; male abdomen filling entire space between coxae of pereiopods 5, with somites 3–5 fused; thoracic sternum wide; sternite 7 with posterolateral prolongation; sternite 8 visible in ventral view, overlying posterior of sternite 7; sterno-abdominal cavity reaching anterior of sternite 4; fingers of chelipeds long, elongate, not dark in color.

Type and sole included genus.

Carinocarcinoides Karasawa and Fudouji, 2000.

Discussion.

Karasawa and Fudouji (2000) originally placed Carinocarcinoides within the Carcinoplacinae. However, the phylogenetic analysis strongly suggests that the genus should not be assigned to the Carcinoplacinae (= Goneplacinae) based upon examination of type and newly obtained specimens. Carinocarcinoides is derived as the sister to the Trogloplacinae and Chasmocarcininae, and lacks diagnostic synapomorphies of both subfamilies. Carinocarcinoides is here treated as the type of a new subfamily. Based upon the present phylogenetic analysis, the Carinocarcinoidinae belongs in a monophyletic group with the Trogloplacinae and Chasmocarcininae within the Goneplacidae.

Fossil records.

Two species have been recorded from the lower Oligocene of Japan (Table 7).

Table 7

Distributions and geologic ranges of recognized fossil species of the subfamily Carinocarcinoidinae.

i1342-8144-7-2-129-t07.gif

Subfamily Trogloplacinae Guinot, 1986

Diagnosis.

Carapace rounded, sometimes poorly calcified; dorsal regions poorly defined; front straight with shallow median indentation; notch between frontal margin and supraorbital angle indistinct; upper orbital margin narrow without fissures; anterolateral margin cristate, entire or toothed; eye stalk short; merus of maxilliped 3 subquadrate, about as long as ischium; male abdomen not filling entire space between coxae of pereiopods 5, with somites 3–5 fused; thoracic sternum wide with sutures all interrupted; sternite 7 with posterolateral prolongation; sternite 8 visible in ventral view, overlying posterior part of sternite 7, with supplementary plate; chelipeds without dark-colored fingers; dactyli of pereiopods 2–5 with or without corneous tips; dactyli of pereiopods 5 styliform with or without setae; gonopod 1 stout, sinuous, with simple apex; gonopod 2 long, about as long as gonopod 1, with flagellum about same length as peduncle (from Davie and Guinot, 1996).

Type genus.

Trogloplax Guinot, 1986.

Genera included.

Australocarcinus Davie, 1987; Trogloplax.

Remarks.

The Trogloplacinae is derived as the sister group to the Chasmocarcininae based upon the present phylogenetic analysis. Davie and Guinot (1996) showed that the subfamily is most closely related to the Chasmocarcininae and is separated from it by the suture of the antennular region and differences in length and shape of the male gonopods.

Fossil records.

None.

Subfamily Chasmocarcininae Serène, 1964 emend.

Diagnosis.

Carapace with poorly defined dorsal regions; front straight with shallow median notch; notch between frontal margin and supraorbital angle indistinct; orbit usually small without upper orbital fissures; anterolateral margin entire or toothed, tapering anteriorly; eye stalk short; merus of maxilliped 3 suboval, about as long as ischium; male abdomen not filling entire space between coxae of pereiopods 5, with somites 3–5 fused; thoracic sternum wide with sutures all interrupted; sternite 7 with posterolateral prolongation; sternite 8 visible in ventral view, overlying posterior part of sternite 7, with supplementary plate; sterno-abdominal cavity reaching anterior of sternite 4; chelipeds without dark-colored fingers; fingers sometimes elongate, deflexed; dactyli of pereiopods 2–5 without corneous tips; dactyli of pereiopods 5 sickleshaped with setae; gonopod 1 stout, sinuous, with simple apex; gonopod 2 long, but shorter than gonopod 1, with flagellum much shorter than peduncle.

Type genus.

Chasmocarcinus Rathbun, 1898.

Genera included.

Camatopsis Alcock and Anderson, 1899, Chasmocarcinus; Chasmocarcinops Alcock, 1900, Collinsius Karasawa, 1993; Falconoplax Van Straelen, 1933, Gillcarcinus Collins and Morris, 1978; Hephthopelta Alcock, 1899; Mioplax Bittner, 1884; Orthakrolophos Schweitzer and Feldmann, 2001a; Parapilumnus Kossmann, 1877; Scalopidia Stimpson, 1858.

Discussion.

Serène (1964) originally included Megaesthesius Rathbun, 1909, within the Chasmocarcininae while Davie and Guinot (1996) excluded this genus from the subfamily. We concur with Davie and Guinot. Ng (2002) moved Parapilumnus, previously assigned to the Pilumnidae, to the Chasmocarcininae.

Schweitzer and Feldmann (2001a) recognized three extinct genera within the Chasmocarcininae. It is here expanded to include two genera Gillcarcinus and Mioplax. Collins and Morris (1978) erected the monotypic genus Gillcarcinus from the middle Eocene of Pakistan and referred the genus to the Xanthidae. The genus is moved to the Chasmocarcininae by having a narrow upper orbital margin without notches, a wide thoracic sternum, a narrow male abdominal somite 3, which does not fill the entire space between pereiopods 5, and fused male abdominal somites 3–5. Glaessner (1969) placed Mioplax from the Miocene of Austria within the Goneplacinae; however, the genus possesses a small orbit and long, slender deflexed fingers of chelipeds. Both characters strongly suggest that Mioplax should be assigned to the Chasmocarcininae. Gillcarcinus has three weakly developed anterolateral spines and Mioplax bears a well developed anterolateral spine. Most members of the Chasmocarcininae lack anterolateral spines, while the extant Hephthopelta aurita Rathbun, 1932, has two sharp anterolateral spines.

Fossil records.

Six genera are known from the fossil record (Table 8). Collinsius, Falconoplax, and Orthakrolophos are extinct genera. Fossil members of Chasmocarcinus are known from the Eocene of Antarctica and Miocene of the U.S.A.

Table 8

Distributions and geologic ranges of recognized fossil species of the subfamily Chasmocarcininae.

i1342-8144-7-2-129-t08.gif

A review of remaining fossil genera

"Pilumnoplax Stimpson, 1858"

Guinot (1969a, b, c; Tucker and Feldmann, 1990) have already discussed the nomenclatural status of the generic name Pilumnoplax, and have shown that the genus was a heterogenous group. Bachmayer (1953b) described a new species, Pilumnoplax carnuntinus, from the Miocene of Austria based upon a single incomplete specimen. In this specimen the front, a part of the upper orbital margin, and a part of the gastric region have been preserved; therefore, the species is not classified within any known genus of the Goneplacidae (Müller, 1984; 1998). Pilumnoplax soledadensis Rathbun, 1926, described from the Eocene of the U.S.A., was moved to the panopeid genus Panopeus H. Milne Edwards, 1834 (Schweitzer, 2000).

Glyphithyreus Reuss, 1859 (= Plagiolophus Bell, 1858 non Pomel, 1847)

Glyphithyreus Reuss, 1859, has been placed within the goneplacid Carcinoplacinae (Balss, 1957, Glaessner, 1969 and many subsequent workers). Glyphithyreus lacks the poorly defined dorsal carapace regions, a straight front margin without median notch, the upper orbital margin with an indistinct supraorbital angle and without fissures, and a wide male abdomen with all free somites, all of which are diagnostic characters of the Goneplacinae (= Carcinoplacinae). Glyphithyreus is here placed in the panopeid Eucratopsinae because the carapace has well defined dorsal regions, the front consists of two rounded lobes, and the narrow male abdomen has fused somites 3–5. Previously known species of Glyphithyreus include: G. ellipticus Bittner, 1875, from the Eocene of Italy; G. markgrafi (Lörenthey, 1907 [1909]), from the Eocene of Egypt; G. sturgeoni Feldmann et al., 1998, from the Eocene of the U.S.A.; G. weaveri (Rathbun, 1926) from the Eocene of the U.S.A.; G. wetherelli (Bell, 1858) (type species) from the Eocene of Europe, Senegal, and Pakistan; and ? G. wichmanni Feldmann et al., 1995, from the Danian of Argentina. Among these, Glyphithyreus weaveri was moved to the euryplacine Orbitoplax (Schweitzer, 2000).

Galenopsis A. Milne Edwards, 1865

A. Milne Edwards (1865) erected the genus Galenopsis containing five species within his “Galénides”. Subsequently, Glaessner (1929) placed the genus within the Xanthidae, Balss (1957) and Glaessner (1969) removed it to the goneplacid Carcinoplacinae, and Schweitzer (2000) reassigned it to the Pilumnidae. We agree with Schweitzer's opinion. In Galenopsis a narrow, deflexed frontal margin is medially interrupted with prominent median lobes, and the narrow upper orbital margin possesses a distinct supraorbital angle. These characters do not match the diagnostic characters of the Goneplacinae (= Carcinoplacinae) but match those of the Pilumnidae as defined by Schweitzer (2000).

Galenopsis contains numerous species (Via, 1969) from the Eocene-Pliocene of Europe, Africa, India, and Fiji (Glaessner, 1969). Among these Müller and Collins (1991a) proposed a new monotypic genus, Lobogalenopsis, for Galenopsis quadrilobatus Lörenthey, 1897, from the upper Eocene of Hungary. Lobogalenopsis was also excluded from the Goneplacidae and has been transferred to the Pilumnidae (Schweitzer, 2000).

Palaeograpsus Bittner, 1875

Palaeograpsus has long been placed within the Grapsidae MacLeay, 1838 (Glaessner, 1969). Previously known species of the genus are Palaeograpsus attenuatus Bittner, 1875, P. bartonensis Quayle and Collins, 1981, P. depressus Quayle and Collins, 1981, P. guerini Via, 1959, P. inflatus Bittner, 1875 (type species), P. loczyanus Lörenthey, 1898a and P. parvus (Müller and Collins, 1991b) from the Eocene of Europe; and P. bittneri Morris and Collins, 1991 from the Pliocene of Brunei. Among these, Schweitzer and Feldmann (2001a) moved three species, P. bartonensis, P. bittneri and P. depressus, to Orthakrolophos within the Goneplacidae. Palaeograpsus guerini is similar to members of Orthakrolophos, but is characterized by having transverse ridges on the dorsal carapace, which are absent in Orthakrolophos; therefore, Schweitzer and Feldmann (2001a) did not include the species in Orthakrolophos.

Via (1969) suggested that P. loczyanus closely resembles members of Carcinoplax within the Goneplacinae. Karasawa and Kato (2001) also suggested that P. inflatus and P. loczyanus possess carapace and cheliped characters most like those of Carcinoplax and moved Palaeograpsus to the Goneplacidae. Reexamination of P. inflatus described by De Angeli (1995) strongly suggests that the genus does not belong within the Goneplacinae but within the panopeid Eucratopsinae or the Pseudorhombilidae because the front has a median notch (De Angeli, 1995, figs. 3.2, 3.4, pl. 2, figs. 2, 4), there are two fissures on the upper orbital margin (De Angeli, 1995, figs. 3.2, 3.4, pl. 2, figs. 2, 4), male abdominal somites 3 and 4 are incompletely fused (De Angeli, 1995, fig. 3.4, pl. 2, fig. 4) and male abdominal somite 1 fills the entire space between the coxae of pereiopods 5 (De Angeli, 1995, pl. 2, figs. 2–4). The panopeid Eucratopsinae is quite similar to the Pseudorhombilidae, and the differentiation between them is mainly based upon the male gonopod morphology (Hendrickx, 1998). However, male abdominal somite 1 in members of the Eucratopsinae does not fill the entire space between coxae of pereiopods 5; it is therefore considered best to place Palaeograpsus inflatus within the Pseudorhombilidae.

The monotypic genus Carinocarcinus Lörenthey, 1898b, described from the middle Eocene of Germany, possesses carapace and cheliped characters like those of Palaeograpsus sensu stricto; however, in Carinocarcinus the carapace is much wider than long with four anterolateral teeth. Carinocarcinus may be also referred to the Pseudorhombilidae.

Telphusograpsus Lörenthey, 1902

Glaessner (1969) showed that Telphusograpsus Lörenthey, 1902, from the Eocene of Rumania, was synonymous with the grapsid genus Varuna H. Milne Edwards, 1830; however, Karasawa and Kato (2001) suggested that Telphusograpsus is an independent genus and probably belongs to the Goneplacidae. Telphusograpsus, represented only by a carapace specimen, has a distinct supraorbital angle and upper orbital fissures; therefore, it could be referred to the Mathildellinae, the Euryplacinae, the Panopeidae or the Pseudorhombilidae. Complete familial and subfamilial arrangements of the genus must await discovery of the thoracic sternum and the male abdomen.

Laevicarcinus Lörenthey in Lörenthey and Beurlen, 1929

Laevicarcinus was originally placed within the Carcinoplacidae (= Goneplacidae), and within the goneplacid Carcinoplacinae by Balss (1957) and Glaessner (1969). Crosnier and Guinot (1969) indicated that the genus has a close resemblance to Platychelonion. Müller and Collins (1991a) reexamined the type specimen of the type species, Laevicarcinus egerensis Lörenthey in Lörenthey and Beurlen, 1929, from the upper Eocene of Hungary and removed Laevicarcinus to the Panopeidae.

Martinezicancer Van Straelen, 1939

Van Straelen (1939) described a new genus and species, Martinezicancer schencki, from the Palaeogene of California and suggested that the genus might represent a new family. Glaessner (1969) placed the genus within the Carcinoplacinae. Martinezicancer schencki has well defined dorsal regions, an arcuate protogastric ridge, an epibranchial region with two ovoid swellings, a mesogastric region with low transverse ridge, and a broad cardiac region with lobate swellings along lateral margins; therefore, it is assigned to Archaeopus Rathbun, 1908, redefined by Schweitzer and Feldmann (2001b), of the Retroplumidae Gill, 1894, and Martinezicancer becomes a junior synonym of Archaeopus.

Carcinoplacoides Kesling, 1958

The monotypic genus, Carcinoplacoides, erected with C. flottei Kesling, 1958, from the Pleistocene of Guam, was placed within the Carcinoplacinae (Kesling, 1958; Glaessner, 1969). Schweitzer et al. (2002) indicated that the species is synonymous with Libystes nitidus A. Milne Edwards, 1867, of the Portunidae.

Styrioplax Glaessner, 1969

Glaessner (1969) gave Styrioplax as a replacement generic name for Microplax Glaessner, 1928. This monotypic genus contains S. exiguus (Glaessner, 1928) from the Miocene of Austria. Glaessner (1969) did not place it in any known subfamily within the Goneplacidae. This genus is characterized by a small-sized carapace, a straight frontal margin with a median notch, small orbits, and a distinctly narrowed male abdomen. These characters indicate that the genus may belong to the Trogloplacinae, Chasmocarcininae, or pilumnid Rhizopinae sensu lato. The familial and subfamilial placements of Styrioplax remain obscure because detailed characters of the male abdomen of S. exiguus are poorly known.

Caprocancer Müller and Collins, 1991a and Eoplax Müller and Collins, 1991a

Müller and Collins (1991a) erected the two new goneplacid genera, Caprocancer and Eoplax, from the upper Eocene of Hungary. However, Müller and Collins (1991a) did not make any comparisons between Caprocancer and any known genera of decapods and indicated that there was a similarity between Eoplax and the grapsid genus Pachygrapsus Randall, 1840. The familial placement of both genera remains obscure because they are based upon poorly preserved carapace specimens.

Chumaoia Hu and Tao, 1996

Hu and Tao (1996) erected a new monotypic genus, Chumaoia, with C. johnferi Hu and Tao, 1996, from the Miocene of Taiwan. Karasawa (1997, p. 67, footnote) showed that the genus is a junior synonym of Typilobus Stoliczka, 1871, of the Leucosiidae Samouelle, 1819.

Pregeryona Hu and Tao, 1996

Hu and Tao (1996) described a new genus and species, Pregeryona taiwanica, from the Miocene of Taiwan. The familial placement of the genus is doubtful because their description was brief and their materials poorly preserved.

Maingrapsus Tessier et al., 1999 and Paracorallicarcinus Tessier et al., 1999

Tessier et al., (1999) erected a new grapsid genus, Maingrapsus, from the Eocene of Italy. Karasawa and Kato (2001) suggested that the genus resembles Paracorallicarcinus Tessier et al., 1999 and moved it to the Goneplacidae. Maingrapsus and Paracorallicarcinus possess carapace characters like those of the extant Georgeoplax Türkay, 1983 of the Pilumnidae (as Goneplacidae by Karasawa and Kato, 2001); therefore, both genera are removed to the Pilumnidae.

The current status of the sixteen genera discussed above is listed in Table 9.

Table 9

The current status of the remaining extinct genera previously assigned to the Goneplacidae.

i1342-8144-7-2-129-t09.gif

Acknowledgments

We thank T. Komai (Natural History Museum and Institute, Chiba), L. Lemaitre and W. C. Blow (National Museum of Natural History, Smithsonian Institution, Washington, D.C.), P. Müller (Hungarian Geological Survey, Hungary), C. Beschin and A. De Angeli (Museo Civico “G. Zannato”, Italy), and T. Sato (Kanagawa Prefectural Museum of Natural History) for loans of their material. We also thank D. Guinot (Muséum national d'Histoire naturelle, Paris), and Z. Stevcic (Center for Marine Research, Croatia) for providing valuable comments about some taxa of the Goneplacidae; T. Kimura (Gunma Prefectural Museum of Natural History) for providing valuable information about the phylogenetic analysis; and A. Garassino (Museo Civico di Storia Naturale in Milano, Italy) and A. De Angeli (Museo Civico “G. Zannato”, Italy) for providing copies of classical literature. We are deeply indebted to C. L. McLay (University of Canterbury, New Zealand), P. K. L. Ng (National University of Singapore), and C. E. Schweitzer (Kent State University, U.S.A.) for reading our manuscript and providing useful comments. Special thanks are due to R. M. Feldmann (Kent State University, U.S.A.) and an anonymous referee for their review of the manuscript. Travel for examination of Hungarian fossils was provided by a special fund from the Mizunami Municipal Government to Karasawa.

References

1.

A. Adams and A. White . 1848. Crustacea. In,. A. Adams The Zoology of the Voyage of H. M. S. Samarang, 1843–1846, viii + 66 p., 13 pls. London. Google Scholar

2.

A. Alcock 1894. Natural history notes from the Indian Marine Survey Steamer “Investigator”. Series II. no. 1On the re- sult of deep-sea dredging during the season of 1890-1891. Annals and Magazine of Natural History, Series 6 vol. 13:p. 400–411. Google Scholar

3.

A. Alcock 1899. An account of the deep-sea Brachyura collected by the Royal Indian Marine Survey Ship 'Investigator'. 86. p. Calcutta. Google Scholar

4.

A. Alcock 1898. The family Xanthidae: the Brachyura Cyclo- metopa, Part I: Materials for a carcinological fauna of India. no. 3:Journal of the Asiatic Society of Bengal vol. 67:p. 67–233. Google Scholar

5.

A. Alcock 1900. Brachyura Catometopa or Grapsoidea: Materials for a carcinological fauna of India. no. 6:Journal of the Asiatic Society of Bengal vol. 69:p. 279–456. Google Scholar

6.

A. Alcock and A.R.S. Anderson . 1899. Natural history notes from H. M. Royal Indian Marine Survey Ship 'Investigator'. H. T and R.N. Heming . commanding. Ser III,. no. 2:An account of the deep-sea Crustacea dredged during the survey season of 1897-1898. Annals and Magazine of Natural History, Series 7 vol. 3:p. 1–27. p. 278–292. Google Scholar

7.

F. Bachmayer 1953a. Goneplax gulderi eine neue Crustaceen- Species aus dem tortonischen Tegel des Wiener Beckens. Paläontologische Zeitschrift vol. 27:nos. 3–4. p. 143–145. Google Scholar

8.

F. Bachmayer 1953b. Die Decapodenfauna des tortonischen Leithakalkes von Deutsch-Alternburg (Niederösterreich). Mitteilungen der Geologischen Gesellschaft in Wien vol. 44:(for 1951). p. 237–262. Google Scholar

9.

H. Balss 1957. Decapoda. In, Dr. H. G. Bronns Klassen und Ordnungen des Tierreichs. Band 5, Arthropoda, Abteilung 1, Buch 7, Lieferung. 12–p. 1505–1672. Google Scholar

10.

T. Bell 1858. A Monograph of the Fossil Malacostracous Crustacea of Great Britain. Part I. Crustacea of the London Clay. 44. p. Palaeontographical Society. London. Google Scholar

11.

A. Bittner 1875. Die Brachyuren des Vicentinischen Tertiärge- birges. Denkschriften der kaiserlichen Akademie der Wissens- chaften in Wien vol. 34:p. 63–103. Google Scholar

12.

A. Bittner 1884. Beiträge zur Kenntnis tertiärer Brachyurenfaunen. Denkschriften der kaiserlichen Akademie der Wissenschaften in Wien vol. 48:p. 15–30. Google Scholar

13.

W.C. Blow and R.H. Bailey . 1992. Chasmocarcinus robertsi a new crab species from the Miocene of Virginia, with notes on the genus Falconoplax (Crustacea, Decapoda, Goneplacidae). Tulane Studies in Geology and Paleontology. vol. 25:p. 175–185. Google Scholar

14.

K. Bremer 1994. Branch support and tree stability. Cladistics vol. 10:p. 295–304. Google Scholar

15.

G. Colosi 1923. Una specie fossile de Gerionide (Decapodi Brachiuri). Bolettino della Societá dei Naturalisti in Napoli vol. 37:p. 248–255. Google Scholar

16.

J.S.H. Collins and S.F. Morris . 1976. Tertiary and Pleistocene crabs from Barbados and Trinidad. Palaeontology vol. 19:p. 107–131. Google Scholar

17.

J.S.H. Collins and S.F. Morris . 1978. New lower Tertiary crabs from Pakistan. Palaeontology vol. 21:p. 957–981. Google Scholar

18.

C. Crema 1895. Sopra alcuni decapodi tertiarii del Piemonte. Atti della Reale Academia di Scienze di Torino vol. 30:p. 664–681. Google Scholar

19.

A.W. Cresswell 1886. Notes on some fossil crabs from the Miocene rocks of Corio Bay. Victorian Naturalist vol. 3:p. 86–91. Google Scholar

20.

A. Crosnier and D. Guinot . 1969. Un nouveau crabe ouest-africain. Platychelonion planissimum gen. nov., sp. nov. Bulletin du Muséum national d'Histoire naturelle, Paris, série 2 vol. 41:p. 725–730. Google Scholar

21.

J.D. Dana 1851. On the classification of the Cancroidea. American Journal of Science and Arts, Series 2 vol. 12:p. 121–131. Google Scholar

22.

J.D. Dana 1852. Crustacea. In, United States Exploring Expedition during the Years 1838, 1839, 1840, 1841, 1842 Under the Command of Charles Wilkes, U.S.N. vol. 13:p. 1–1620. Google Scholar

23.

P.J.F. Davie 1987. A new genus and species of goneplacid (Crustacea: Brachyura) from Queensland, Australia. Memoirs of the Queensland Museum vol. 25/no. 2:p. 259–264. Google Scholar

24.

P.J.F. Davie 1989. A re-appraisal of. Heteropanope Stimpson, and Pilumnopeus A. Milne Edwards (Crustacea: Decapoda: Pilumnidae) with descriptions of new species and new genera. Memoirs of the Queensland Museumvol. 27:p. 129–156. Google Scholar

25.

P.J.F. Davie and D. Guinot . 1996. Two new freshwater crabs in. Australocarcinus Davie, with remarks on Trogloplacinae Guinot and Goneplacidae MacLeay (Crustacea: Decapoda: Brachyura). Memoirs of the Queensland Museum vol. 39/no. 2:p. 277–287. Google Scholar

26.

A. De Angeli 1995. Crostacei dell'Eocene superiore di “Fontanella” di Grancona (Vicenza-Italia settentrionale). Studi e Ricerche, Associazione Amici del Museo Civico “G. Zannato” Montecchio Maggiore (VI) p. 7–24. Google Scholar

27.

W. De Haan 1833-1850. Crustacea. In Siebold P.Siebold von. ed., Fauna Japonica sive descriptio animalium, quae in Itinere per Japoniam, Jussu et auspiciis superiorum, qui Summun in India Batava Imperium tenent, suscepto, annis 1823–1830 collegit, notis, observationibus et adumbrationibus illustravit. p. i–xvii + i–xxxi + ix–xvi + 1–243, pls. A–J + L–Q + 1–55. Lugduni-Batavorum.  Google Scholar

28.

F. Doflein 1904. Brachyura. In, Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer “Valdivia” 1898-1899 xiv + 314 p. Jena. Google Scholar

29.

T. Eriksson 1999. AutoDecay ver. 4.0. Bergius Foundation, Royal Swedish Academy of Sciences. Stockholm. Google Scholar

30.

J.C. Fabricius 1793. Entomologia Systematica Emendata et Aucta. Secundum Classes, Ordines, Genera, Species, Adjectis Synonimis, Locis, Observationibus, Descriptionibus. vol. 2.viii + 519 p. Hafniae. Google Scholar

31.

D.L. Felder and N.N. Rabalais . 1986. The genera. Chasmocarcinus Rathbun and Speocarcinus Stimpson on the Continental Shelf of the Gulf of Mexico, with descriptions of two new species (Decapoda: Brachyura: Goneplacidae). Journal of Crustacean Biologyvol. 6:p. 547–575. Google Scholar

32.

R.M. Feldmann, K.L. Bice, C. Schweitzer Hopkins, E.W. Salva, and K. Pickford . 1998. Decapod crustaceans from the Eocene Castle Hayne Limestone, North Carolina: Paleoceanographic implications. The Paleontological Society Memoir 48 (Supplement to Journal of Paleontologyvol. 72:) p. 1–28. Google Scholar

33.

R.M. Feldmann, Casadío, L. Chirino-Galvez, and M. Aguirre-Urreta . 1995. Fossil decapod crustaceans from the Jaguel and Roca Formations (Maastrichtian-Danian) of the Neuquén basin, Argentina. The Paleontological Society Memoir 43 (Supple- ment to Journal of Paleontologyvol. 69:) p. 1–22. Google Scholar

34.

R.M. Feldmann and I.W. Keyes . 1992. Systematic and stratigraphic review with catalogue and locality index of the Mesozoic and Cenozoic decapod Crustacea of New Zealand. New Zealand Geological Survey Record no. 45:p. 1–73. Google Scholar

35.

R.M. Feldmann and P.A. Maxwell . 1990. Late Eocene decapod Crustacea from North Westland, South Island, New Zealand. Journal of Paleontology vol. 64:p. 779–797. Google Scholar

36.

R.M. Feldmann and W.J. Zinsmeister . 1984. New fossil crabs (Decapoda: Brachyura) from the La Meseta Formation (Eocene) of Antarctica: Paleogeographic and biogeographic implications. Journal of Paleontology vol. 58:p. 1046–1061. Google Scholar

37.

J. Felsenstein 1985. Confidence limits of phylogenies: an approach using the bootstrap. Evolution vol. 39:p. 783–791. Google Scholar

38.

H. Filhol 1885. Catalogue des crustacés de la Nouvelle-Zélande, des îles Auckland et Campbell. In, Mission de l'île Campbell, Recueil de mémoires, rapports, et documents relatifs à l'obser- vation du passage de Vénus sur le soleil vol. 3:p. 349–510. Google Scholar

39.

T. Gill 1894. A new bassalian type of crab. American Naturalist vol. 28:p. 1043–1045. Google Scholar

40.

M.F. Glaessner 1928. Die Dekapodenfauna des österreichischen Jungtertiärs. Jahrbuch der Geologischen Bundesanstalt vol. 78:p. 161–219. Google Scholar

41.

M.F. Glaessner 1929. Crustacea Decapoda. In F.Pompeckji Pompeckji , editor. ed., Fossilium Catalogus 1. Animalia 464 p. Berlin. Google Scholar

42.

M.F. Glaessner 1960. The fossil decapod Crustacea of New Zealand and the evolution of the order Decapoda. New Zealand Geological Survey Palaeontological Bulletin vol. 31:p. 1–63. Google Scholar

43.

M.F. Glaessner 1969. Decapoda. In. R.Moore Moore ed., Treatise on Invertebrate Paleontology, Part R, Arthropoda 4. p. R399–R533. R626–R628. Geological Society of America and University of Kansas Press. Google Scholar

44.

D. Guinot 1969a. Les Goneplacidae: Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures, VII. Bulletin du Muséum national d'Histoire naturelle, Paris, série 2 vol. 41:p. 241–265. Google Scholar

45.

D. Guinot 1969b. Les Goneplacidae (suite): Recherches prélimi- naires sur les groupements naturels chez les Crustacés Décapodes Brachyoures, VII. Bulletin du Muséum national d'Histoire naturelle, Paris, série 2 vol. 41:p. 507–528. Google Scholar

46.

D. Guinot 1969c. Les Goneplacidae (suite): Recherches prélimi- naires sur les groupements naturels chez les Crustacés Décap odes Brachyoures, VII. Bulletin du Muséum national d' Histoire naturelle, Paris, série 2 vol. 41:p. 688–724. Google Scholar

47.

D. Guinot 1970 [1971]. Synthèse et bibliographie: Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures, VIII. Bulletin du Muséum national d'Histoire naturelle, Paris, série 2 vol. 42:p. 1063–1090. Google Scholar

48.

D. Guinot 1978. Principes d'une classification évolutive des Crustacés Décapodes Brachyoures. Bulletin Biologique de le France et de le Belgique vol. 112:p. 211–292. Google Scholar

49.

D. Guinot 1985. Crabes bathyaux de l'île de La Réunion; descrip- tion de Cyrtomaia guillei sp. nov., de Platypilumnus inermis; sp. nov. et de Psopheticus vocans sp. nov. (Crustacea Decapoda, Brachyura). Comité National Français de la Recherche Antarctique vol. 55:p. 7–31. Google Scholar

50.

D. Guinot 1986. Description d'un crabe cavernicole aveugle de Nouvelle-Bretagne (Papouasie Nouvelle-Guinée), Trogloplax joliverti gen. nov. sp. nov., et éstablissement d'une sous- famille nouvelle, Trogloplacinae subfam. nov Comptes Rendus de L'Académie des Sciences, Paris, séries 3 vol. 303:p. 307–312. Google Scholar

51.

D. Guinot 1989. Le genre. Carcinoplax H. Milne Edwards, 1852 (Crustacea, Brachyura: Goneplacidae). In Crosnier, A. ed., Résultats des Campagnes MUSORSTOM, tome 5, Mémoires du Muséum national d'Histoire naturelle, série A vol. 144:p. 265–345. Google Scholar

52.

D. Guinot 1990. Crustacea Decapoda: Le genre. Psopheticus Wood-Mason, 1892 (Goneplacidae). In Crosnier A. ed., Résultats des Campagnes MUSORSTOM, tome 6. Mémoires du Muséum national d'Histoire naturelle, série A vol. 145:p. 331–367. Google Scholar

53.

D. Guinot and B. Richer de Forges . 1981. Crabes de profondeur, nouveaux ou rares, de I'Indo-Pacifique (Crustacea, Decapoda, Brachyura) (Deuxième partie). Bulletin du Muséum national d'Histoire naturelle, Paris, 4ème série, 3, A vol. 4:p. 227–260. Google Scholar

54.

C. Heller 1861. Beiträge zur Crustaceen-Fauna des rothen Meeres. Sitzungsberichte der mathematisch-naturwissenschaftlischen Classe der Kaiserlichen Akademie der Wissenschaften vol. 43:p. 297–374. Google Scholar

55.

M.E. Hendrickx 1998. A new genus and species of “goneplacid- like” brachyuran crab (Crustacea: Decapoda) from the Gulf of California, Mexico, and a proposal for the use of the family Pseudorhombilidae Alcock, 1900. Proceedings of the Biologi- cal Society of Washington vol. 111:p. 634–644. Google Scholar

56.

P.-W. Hsueh and J.-F. Huang . 2002. Crabs of the family Goneplacidae (Decapoda, Brachyura) from Taiwan. Crusta- ceana vol. 75:p. 111–136. Google Scholar

57.

C.-H Hu and H.-J. Tao . 1985. Tertiary crustacean fossils from Taiwan. Proceedings of the Geological Society of China vol. 31., 239–260.  Google Scholar

58.

C.-H Hu and H.-J. Tao . 1996. Crustacean Fossils of Taiwan. 228. p. Taipei, Taiwan. Google Scholar

59.

H. Ikeda 1998. The Deep-sea Crabs of Sagami Bay. 180. p. Hayama Shiosai Museum. Kanagawa. Google Scholar

60.

R. Imaizumi 1961. A critical review and systematic descriptions of known and new species of. Carcinoplax from Japan. Science Reports of the Tohoku University, Series 2 (Geology) vol. 32:p. 155–193. Google Scholar

61.

R.J.F. Jenkins 1975. The fossil crab. Ommatocarcinus corioensis (Cresswell) and a review of related Australian species. Memoirs of the National Museum of Victoria vol. 36:p. 33–67. Google Scholar

62.

H. Karasawa 1993. Cenozoic decapod Crustacea from southwest Japan. Bulletin of the Mizunami Fossil Museum no. 20:p. 1–92. Google Scholar

63.

H. Karasawa 1997. A monograph of Cenozoic stomatopod, decapod, isopod and amphipod Crustacea from west Japan. Monograph of the Mizunami Fossil Museum no. 8:p. 1–81. Google Scholar

64.

H. Karasawa and Y. Fudouji . 2000. Palaeogene decapod Crustacea from the Kishima and Okinoshima Groups, Kyushu, Japan. Paleontological Research vol. 4/no. 4:p. 239–253. Google Scholar

65.

H. Karasawa and K. Inoue . 1992. Decapod crustaceans from the Miocene Kukinaga Group, Tanegashima Island, Kyushu, Japan. Tertiary Research vol. 14:p. 73–96. Google Scholar

66.

H. Karasawa and H. Kato . 2001. The systematic status of the genus. Miosesarma Karasawa, 1989 with a phylogenetic analysis within the family Grapsidae and a review of fossil records (Crustacea: Decapoda: Brachyura). Paleontological Research vol. 5/no. 4:p. 259–275. Google Scholar

67.

H. Kato 1996. Miocene decapod Crustacea from the Chichibu Basin, Central Japan. Transactions and Proceedings of the Palaeontological Society of Japan, New Series no. 183:p. 500–521. Google Scholar

68.

R.V. Kesling 1958. Fossil crabs from Guam. Contributions from the Museum of Paleontology, University of Michigan vol. 14:p. 207–263. Google Scholar

69.

J.R. Kinaham 1856. Remarks on the habits and distribution of ma- rine Crustacea on the eastern shores of Port Philip, Victoria, Australia, with descriptions of undescribed species and genera. Journal of the Royal Dublin Society vol. 1:p. 111–134. Google Scholar

70.

R. Kossmann 1877. Malacostraca (1: Theil Brachyura). In Kossmann R. ed., Zoologische Ergebnisse einer Reise in die Küstengebiete des Roten Meeres, Erste Hälfte III. p. 1–66.Leipzig. Google Scholar

71.

P.A. Latreille 1802-1803. Histoire naturelle, générale et parti- culière, des crustacés et des insectes.. vol. 3:467. p. F. Dufart. Paris. Google Scholar

72.

W.E. Leach 1814. Crustaceology. In D. Brewster, The Edinburgh Encyclopaedia vol. 7:p. 383–437. Google Scholar

73.

W.E. Leach 1815. A tabular view of the external characters of four classes of animals, which Linné arranged under Insecta, with the distribution of the genera composing three of these classes into orders, and description of several new genera and species. Transactions of the Linnean Society of London vol. 11., 306– 400.  Google Scholar

74.

W.E. Leach 1817-1818. A general notice of the animals taken by Mr. John Cranch, during the expedition to explore the source of the River Zaire. In. J.K. Tuckey Narrative of an Expedition to Explore the River Zaire, Usually Called the Congo, in South Africa, in 1816, under the Direction of Captain J.K. Tuckey R.N. To Which Is Added, the Journal of Professor Smith, Some General Observations on the Country and its Inhabitants, and an Appendix: Containing the Natural History of that Part of the Kingdom of Congo Through Which the Zaire Flows.. p. 407–419.London. Google Scholar

75.

R. Lemaitre, J. García-Gómez, R Von Sternberg, and N.H. Campos . 2001. A new genus and a new species of crab of family Goneplacidae MacLeay, 1838 (Crustacea: Decapoda: Brachyura) from the tropical western Pacific. Proceedings of the Biological Society of Washington vol. 114:p. 951–963. Google Scholar

76.

C. Linnaeus 1758. Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characte- ribus, Differentiis, Synonymis, Locis, Tomus 1. volume 1, iii + 823 p. Hafmiae. Google Scholar

77.

E. Lörenthey 1897. Adatok Magyarorszåag harmadkorú rákfauná- jához. Mathematikai és Természettudományi értesitö vol. 15:p. 149–169. Google Scholar

78.

E. Lörenthey 1898a. Beiträge zur Decapodenfauna des ungarischen Tertiärs. Természetrajzi Füzetek vol. 21:p. 1–133. Google Scholar

79.

E. Lörenthey 1898b. Ueber die Brachyuren der paläontologischen Sammlung des bayerischen Staates. Természetrajzi Füzetek vol. 21:p. 134–152. Google Scholar

80.

E Lörenthey 1902. Neuere Beiträge zur tertiären Decapodenfauna Ungarns. Mathematische und naturwissenschaftliche Berichte aus Ungarn vol. 18:p. 98–120. Google Scholar

81.

E. Lörenthey 1907 [1909]. Beiträge zur Kenntnis der eozänen Decapodenfauna Aegyptens. Mathematische und naturwissen- schaftliche Berichte aus Ungarn vol. 25:p. 106–152. Google Scholar

82.

E. Lörenthey and K. Beurlen . 1929. Die fossilen Dekapoden der Länder der ungarischen Krone. Geologica Hungarica, Series Palaeontologica part 3, 420 p. Google Scholar

83.

W.S. MacLeay 1838. On the brachyurous decapod Crustacea brought from the Cape by Dr. Smith. In, Illustrations of the Annulosa of South Africa; being a Portion of the Objects of Natural History Chiefly Collected during an Expedition into the Interior of South Africa, under the Direction of Dr. Andrew Smith, in the Years 1834, 1835, and 1836; Fitted out by “The Cape of Good Hope Association for Exploring Central Africa”. p. 53–71.London. Google Scholar

84.

W.P. Maddison and D.L. Maddison . 2001. MacClade Version 4.03 Sinauer Associates, Sunderland, Massachusetts. Google Scholar

85.

R.B. Manning and L.B. Holthuis . 1981. West African Brachyuran crabs. Smithsonian Contributions to Zoology no. 306:p. 1–379. Google Scholar

86.

R.B. Manning and L.B. Holthuis . 1989. Two new genera and nine new species of geryonid crabs (Crustacea, Decapoda, Geryonidae). Proceedings of the Biological Society of Washington vol. 102:p. 50–77. Google Scholar

87.

J.W. Martin 1984. Notes and bibliography on the larvae of xanthid crabs, with a key to the known xanthid zoeas of the western Atlantic and Gulf of Mexico. Bulletin of Marine Science vol. 34:p. 220–229. Google Scholar

88.

J.W. Martin and L.G. Abele . 1986. Notes on male gonopod mor- phology in the brachyuran crab family Panopeidae Ortmann, 1893. sensu Guinot (1978) (Decapoda). Crustaceana vol. 50:p. 182–198. Google Scholar

89.

J.W. Martin and G.B. Davis . 2001. An updated classification of the recent Crustacea. Natural History Museum of Los Angeles County, Science Series no. 39:p. 1–124. Google Scholar

90.

E.J. Miers 1879. On a collection of Crustacea made by Capt. H.C. St. John R. John in the Corean and Japanese Seas. Part. 1. Podophthalmia. With an Appendix By Capt. H. C. St. John. Proceedings of the Zoological Society of London vol. 1879:p. 18–61. Google Scholar

91.

E.J. Miers 1886. Report on the Brachyura collected by H. M. S. Challenger during the Years 1873–1876. Report on the Scientific Results of the H. M. S. Challenger during the Years 1873-1876, Zoology vol. 17.p. xli + 362. Google Scholar

92.

A. Milne Edwards 1865. Monographie des crustacés de la famille cancériens Suite. Annales des Sciences Naturelles, Zoologie, série 4 vol. 20:p. 273–324. Google Scholar

93.

A. Milne Edwards 1867. Descriptions de quelques espèces nouvelles de Crustacés Brachyures. Annales de la Societé entomologique de France vol. 4:p. 263–288. Google Scholar

94.

A. Milne Edwards 1880. études préliminaires sur les Crustacés, lère Partie In Reports on the Results of Dredging under the Supervision of Alexander Agassiz, in the Gulf of Mexico, and in the Caribbean Sea, 1877, 1878, 1879, by the U. S. Coast Survey Steamer “Blake”, Lieut.-Commander C.D. Sigsbee U.S.N. and Commander J.R. Bartlett U.S.N. Commanding. Bulletin of the Museum of Comparative Zoölogy at Harvard College vol. 8:p. 1–68. Google Scholar

95.

H. Milne Edwards 1830. Varuna. In, Dictionnaire classique d'histoire naturelle. vol. 16.p. 511. Paris. Google Scholar

96.

H. Milne Edwards 1834. Histoire naturelle des Crustacés, comprenant l'anatomie, la physiologie et la classification de ces animaux. vol. 1.xxxvs + 468 p. Paris. Google Scholar

97.

H. Milne Edwards 1852. De la famille des ocypodides (Ocypodidae). Second Mémoire. Observations sur les affinités zoologiques et la classification naturelle des crustacés. Annales des Sciences Naturelles, Zoologie, série 3 (Zoology) vol. 18:p. 109–166. Google Scholar

98.

S.F. Morris and J.S.H. Collins . 1991. Neogene crabs from Brunei, Sabah and Sarawak. Bulletin of the British Museum (Natural History), London, (Geology) vol. 47:p. 1–33. Google Scholar

99.

P. Müller 1984. Decapod Crustacea from the Badenian. Geologica Hungarica, Series Palaeontologica vol. 42:p. 1–317. Google Scholar

100.

P. Müller 1998. Heft VII b/2b Crustacea Decapoda. In Flügel, H. W. ed., Catalogus Fossilium Austriae 55 p. Google Scholar

101.

P. Müller and J.S.H. Collins . 1991a. Late Eocene coral-associated decapods (Crustacea) from Hungary. Contributions to Tertiary and Quaternary Geology vol. 28:p. 47–92. Google Scholar

102.

P. Müller and J.S.H. Collins . 1991b. Palaeograpsus parvus (Crustacea, Decapoda), a replacement name for Palaeograpsus bittneri Müller & Collins, 1991, non Palaeograpsus bittneri Morris & Collins, 1991. Contributions to Tertiary and Quaternary Geology vol. 28:p. 140. Google Scholar

103.

P.K.L. Ng 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 Zoology vol. 4:p. 69–111. Google Scholar

104.

P.K.L. Ng 1998. Crabs. In Carpenter, K. E. and Niem, V. H. eds., The living marine resources of the western Central Pacific. Volume 2. Cephalopods, crustaceans, holothurians, and sharks.. p. 1046–1155.Food and Agriculture Organization of the United Nations. Rome. Google Scholar

105.

P.K.L. Ng 2002. The Indo-Pacific Pilumnidae XVI. On the iden- tity 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 vol. 34:p. 209–226. Google Scholar

106.

P.K.L. Ng and L.Liao Liao . 2002. On a new species of. Euryozius Miers, 1886 (Crustacea: Decapoda: Brachyura: Pseudoziidae) from the Philippines with notes on the taxonomy of the genus Proceedings of the Biological Society of Washington. vol. 115:p. 585–593. Google Scholar

107.

P.K.L. Ng and C.-H. Wang . 1994. Notes on the enigmatic genus. Pseudozius Dana, 1851 (Crustacea, Decapoda, Brachyura). Journal of the Taiwan Museum vol. 47:p. 83–99. Google Scholar

108.

P.K.L. Ng, C.-H. Wang, P.-H Ho, and H.-T. Shih . 2001. An annotated checklist of brachyuran crabs from Taiwan (Crustacea: Decapoda). National Taiwan Museum Special Publication Series no. 11:86 p. Google Scholar

109.

A. Ortmann 1893. Die Decapoden-Krebse des Strassburger Museums, VII. Theil. Abtheilung: Brachyura (Brachyura genuina Boas). II. Unterabtheilung: Cancroidea, 1. Gruppe: Cyclometopa. Zoologische Jahrbücher vol. 7:p. 411–495. Google Scholar

110.

A. Pomel 1847. Note critique sur les caractères et les limites du genre Palaeotherium. Archives des Sciences Physique et Naturelle, Genève vol. 5:p. 200–207. Google Scholar

111.

J. Poupin 1996. Atlas des Crustacés Marins Profonds de Polynésie Française. 59. p. Service Mixte de Surveillance Radiologique et Biologique SMSRB, Montlhéry Cedex. France. Google Scholar

112.

W.J. Quayle and J.S.H. Collins . 1981. New Eocene crabs from the Hampshire Basin. Palaeontology vol. 24:p. 733–758. Google Scholar

113.

J.W. Randall 1840. Catalogue of the Crustacea brought by Thomas Nuttall and J.K. Townsend, from the West Coast of North America and the Sandwich Islands, with descriptions of such species as are apparently new, among which are included spe- cies of different localities, previously existing in the collection of the Academy. Journal of the Academy of Natural Sciences of Philadelphia vol. 8:p. 106–147. Google Scholar

114.

S.C.S. Rafinesque 1815. Analyse de la nature, ou tableau de .pb1 l'univers et des corps organisés 224 p. Palermo.  Google Scholar

115.

M.J. Rathbun 1898. The Brachyura of the biological expedition to the Florida Keys and the Bahamas in 1893. Bulletin from the Laboratories of Natural History of the State University of Iowa vol. 4:p. 250–294. Google Scholar

116.

M.J. Rathbun 1908. Descriptions of the fossil crabs from California. Proceedings of the United States National Museum vol. 35:p. 341–349. Google Scholar

117.

M.J. Rathbun 1909. New crabs from the Gulf of Siam. Proceedings of the Biological Society of Washington vol. 22:p. 107–114. Google Scholar

118.

M.J. Rathbun 1914. A new genus and some new species of crabs of the family Goneplacidae. Proceedings of the United States National Museum vol. 48:p. 137–154. Google Scholar

119.

M.J. Rathbun 1916. Description of a new genus and species of fos- sil crab from Port Townsend, Washington. American Journal of Science vol. 41:p. 344–346. Google Scholar

120.

M.J. Rathbun 1918. The grapsid crabs of America. Bulletin of the United States National Museum no. 97:xxii + 461 p., pls. 1– 161. Google Scholar

121.

M.J. Rathbun 1919. West Indian Tertiary decapod crustaceans. Carnegie Institution of Washington Publication no. 291:p. 159–184. Google Scholar

122.

M.J. Rathbun 1926. The fossil stalk-eyed Crustacea of the Pacific slope of North America. Bulletin of the United States National Museum no. 138:I-VII + 155 p., 39 pls. Google Scholar

123.

M.J. Rathbun 1932. Preliminary descriptions of new species of Japanese crabs. Proceedings of the Biological Society of Washington vol. 45:p. 28–38. Google Scholar

124.

J.-M. Remy and F. Tessier . 1954. Décapodes nouveaux de la partie ouest de Sénégal. Bulletin de la Societé Géologique de France, série 6 vol. 4:p. 185–191. Google Scholar

125.

R. Reuss 1859. Zur Kenntnis fossiler Krabben. Denkschriften der Kais. Akademie der Wissenschaften in Wien vol. 17:p. 1–59. Google Scholar

126.

A.L. Rice 1980. Crab zoeal morphology and its bearing on the classification of the Brachyura. Transactions of the Zoological Society of London vol. 35:p. 271–424. Google Scholar

127.

B. Richer de Forges 1996. The genus. Platypilumnus Alcock and description of P. jamiesoni n. sp. from New Caledonia (Crustacea, Decapoda, Brachyura). Records of the Australian Museum vol. 48:p. 1–6. Google Scholar

128.

G. Ristori 1886. I crostacei brachiuri e anomuri del Pliocene Italiano. Bollettino della Società Italiana vol. 5:p. 93–119. Google Scholar

129.

G. Ristori 1889. Un nuovo crostaceo fossile del Giappone. Atti della Società Toscana di Scienze Naturali vol. 7:p. 4–6. Google Scholar

130.

T. Sakai 1969. Two new genera and twenty-two new species of crabs from Japan. Proceedings of the Biological Society of Washington vol. 82:p. 243–280. Google Scholar

131.

T. Sakai 1974. Notes from the carcinological fauna of Japan (V). Researches on Crustacea no. 6:p. 86–112. Google Scholar

132.

T. Sakai 1976. Crabs of Japan and the Adjacent Seas. xxix + 773 p (English volume), 461 p. (Japanese volume), 251 pls. (Plate volume). Kodansha. Tokyo. Google Scholar

133.

T. Sakai 1980. Decapod Crustacea from the Emperor Seamount Chain. Researches on Crustacea supplement. no. 8:p. 1–39. Google Scholar

134.

G. Samouelle 1819. The Entomologist's Useful Compendium, or an Introduction to the Knowledge of British Insects. 486. p. London. Google Scholar

135.

C.E. Schweitzer 2000. Tertiary Xanthoidea (Crustacea: Decapoda: Brachyura) from the west coast of North America. Journal of Crustacean Biology vol. 20:p. 715–742. Google Scholar

136.

C.E. Schweitzer and R.M. Feldmann . 2001a. Differentiation of the fossil Hexapodidae Miers, 1886 (Decapoda: Brachyura) from similar forms. Journal of Paleontology vol. 75:p. 330–345. Google Scholar

137.

C.E. Schweitzer and R.M. Feldmann . 2001b. New Cretaceous and Tertiary decapod crustaceans from western North America. Bulletin of the Mizunami Fossil Museum no. 28:p. 173–210. Google Scholar

138.

C.E. Schweitzer, R.M. Feldmann, A.B. Tucker, and R.E. Belglund . 2000. Eocene decapod crustaceans from Pulali Point, Washington. Annals of Carnegie Museum vol. 69:p. 27–67. Google Scholar

139.

C.E. Schweitzer, P.R. Scott-Smith, and P.K.L. Ng . 2002. New oc- currences of fossil decapod crustaceans (Thalassinidea, Brachyura) from late Pleistocene deposits of Guam, United States Territory. Bulletin of the Mizunami Fossil Museum no. 29:p. 25–49. Google Scholar

140.

R. Serène 1964. Redescription du genre. Megaesthesius Rathbun et definition des Chasmocarcininae, nouvelle sous-famille des Goneplacidae (Decapoda, Brachyura). Crustaceana vol. 7:p. 175–187. Google Scholar

141.

R. Serène 1984. Crustacés Décapodes Brachyoures de l'Ocean Indien occidental et de la Mer Rouge. Xanthoidea: Xanthidae et Trapeziidae. Addendum: Carpiliidae et Menippidae par Alain Crosnier. Faune Tropicale vol. 24., 399 p., 48 pls.  Google Scholar

142.

R Serène and C.L. Soh . 1976. Brachyura collected during the Thai-Danish Expedition (1966). Phuket Marine Biological Center, Research Bulletin no. 12:p. 1–37. Google Scholar

143.

H.B. Stenzel 1944. A new Paleogene catometope crab from Texas. Tehuacana tehuacana. Journal of Paleontology vol. 18:p. 546–549. Google Scholar

144.

W. Stimpson 1858. Prodromus descriptionis animalium evertebratorum, quae in Expeditione ad Oceanum Pacificum Septentrionalem, a Republica Federata missa, Cadwaladaro Ringgold et Johanne Rodgers Ducibus, observavit et descripsit W. Stimpson. Pars. V. Crustacea Ocypodoidea. Proceedings of the Academy of Natural Sciences of Philadelphia vol. 10:p. 93–110. Google Scholar

145.

W. Stimpson 1859. Notes on North American Crustacea. No. I. Annals of the Lyceum of Natural History of New York vol. 7:p. 49–93. Google Scholar

146.

W. Stimpson 1860. Notes on North American Crustacea, in the Museum of the Smithsonian Institution, No. II. Annals of the Lyceum of Natural History of New York vol. 7:p. 177–246. Google Scholar

147.

W. Stimpson 1871. Preliminary report on the Crustacea dredged in the Gulf Stream in the Straits of Florida, by L. F. de Pourtales, Assist. U. S. Coast Survey. Bulletin of the Museum of Comparative Zoölogy at Harvard College vol. 2:p. 109–160. Google Scholar

148.

F. Stoliczka 1871. Observations on fossil crabs from Tertiary de- posits in Sind and Kutch. Memoirs of the Geological Survey of India. Palaeontologia India, Series 7 vol. 14:16. p.  Google Scholar

149.

D.L. Swofford 1999. PAUP*. Phylogenetic Analysis Using Parsimony, Version 4b. Illinois Natural History Survey Champaign, Illinois. Google Scholar

150.

M. Takeda 1976. Studies on the Crustacea Brachyura of the Palau Islands, III. Xanthidae (1). Researches on Crustacea no. 7:p. 69–99. Google Scholar

151.

M. Takeda and S. Miyake . 1969. A small collection of crabs from New Zealand. OHMU (Occasional Papers of Zoological Laboratory, Faculty of Agriculture, Kyushu University ), vol. 2:p. 157–193. Google Scholar

152.

G. Tessier, C. Beschin, A. Busuluni, and A. De Angeli . 1999. Nuovi Brachiuri Eocenici nella cava “Main” di Arzignano (Vicenza-Italia settentrionale). Lavori-Società Veneziana di Scienze Naturali vol. 24:p. 93–105. Google Scholar

153.

A.B. Tucker and R.M. Feldmann . 1990. Fossil decapod crusta- ceans from the lower Tertiary of the Prince William Sound re- .pb2 gion, Gulf of Alaska. Journal of Paleontology vol. 64:p. 409–427. Google Scholar

154.

M. Türkay 1975. Zur Kenntnis der Gattung. Euchirograpsus mit Bemerkungen zu Brachygrapsus und Litocheira (Crustacea: Decapoda). Senckenbergiana Biologica vol. 56:p. 103–132. Google Scholar

155.

M. Türkay 1983. Georgeoplax new genus for Lithocheira glabra Baker, 1906 (Crustacea: Decapoda: Brachyura). Australian Museum Memoir. no. 18:p. 101–105. Google Scholar

156.

V. Van Straelen 1933. Sur des Crustacés Décapodes Cénozoïques du Venezuela. Bulletin du Museé Royal d'Histoire Naturelle de Belgique vol. 9:p. 1–14. Google Scholar

157.

V. Van Straelen 1939. Martinezicancer schencki brachyure nou- veau du Paléocène du Californie. Bulletin du Museé Royal d'Histoire Naturelle de Belgique vol. 15:p. 1–3. Google Scholar

158.

F.J. Vega, T. Cosma, M.A. Coutiño, R.M. Feldmann, T.G. Nyborg, C.E. Schweitzer, and D.A. Waugh . 2001. New middle Eocene decapods (Crustacea) from Chiapas, México. Journal of Paleontology vol. 75:p. 929–946. Google Scholar

159.

L. Via 1959. Décapodos fósiles del Eoceno español. Boletín del Instituto geológico y minero española vol. 70:p. 331–402. Google Scholar

160.

L. Via 1969. Crustáceos decápodos del Eoceno español. Pirineos nos. 91–94, 479 p.  Google Scholar

161.

R. Von Sternberg and N. Cumberlidge . 2001. Notes on the position of the freshwater crabs within the brachyrhynchan Eubrachyura (Crustacea: Decapoda: Brachyura). Hydrobiologia vol. 449:p. 21–39. Google Scholar

162.

A. White 1852. Descriptions of some new species of Annulosa. In Macgillivray J. ed., Narrative of the Voyage of H.M.S. 'Rattlesnake' during the Years 1846-1850. vol. 2:p. 387–395.Boone London. Google Scholar

163.

A. Williams 1984. Shrimps, Lobsters, and Crabs of the Atlantic Coast of the Eastern United States, Maine to Florida. 550. p. Washington, D.C. Google Scholar

164.

J. Wood-Mason 1892. Crustacea. Part I. Illustrations of the Zoology of the Royal Indian Marine Surveying Steamer “Investigator”. pls. 1–5. Calcutta. Google Scholar

165.

M. Yokoyama 1911. Some Tertiary fossils from the Miike coal- field. Journal of the College of Science, Imperial University of Tokyo vol. 27:p. 1–16. Google Scholar
Hiroaki Karasawa and Hisayoshi Kato "The family Goneplacidae (Crustacea: Decapoda: Brachyura): systematics, phylogeny, and fossil records," Paleontological Research 7(2), 129-151, (30 June 2003). https://doi.org/10.2517/prpsj.7.129
Received: 12 June 2002; Accepted: 1 December 2002; Published: 30 June 2003
JOURNAL ARTICLE
23 PAGES


SHARE
ARTICLE IMPACT
Back to Top