Records of brachyuran crabs from the Pliocene (Piacenzian) of Reggio Emilia (Emilia Romagna, N Italy)
Registros de cangrejos braquiuros del Plioceno (Piacenziano) de Reggio Emilia (Emilia Romaña, N Italia)
Giovanni Pasini1, Alessandro Garassino2,*
1 Via Alessandro Volta 16, I-22070 Appiano Gentile (Como), Italia.
]]> 2 Museo di Storia Naturale, Sezione di Paleontologia, Corso Venezia 55, I-20121 Milano, Italia. *alegarassino@gmail.com
Manuscript received: September 20, 2012.
Corrected manuscript received: November 19, 2012.
Manuscript accepted: November 20, 2012.
Abstract
Four species of brachyuran crabs are recorded from the Pliocene (Piacenzian) clays of the Apennine Mountains, all collected in natural gullies around the Castellarano and Monticelli di Quattro Castella, Reggio Emilia (Emilia Romagna, N Italy). These species are already known in the fossil record of Italy, and are assigned to Calappa granulata (Linnaeus, 1758) (Calappidae De Haan, 1833); Chlinocephalus demissifrons Ristori, 1886 (Euryplacidae Stimpson, 1871); Goneplax rhomboides (Linnaeus, 1758) (Goneplacidae MacLeay, 1838); Monodaeus bortolottii Delle Cave, 1988 (Xanthidae MacLeay, 1838). Chlinocephalus demissifrons and Monodaeus bortolottii are reported for the first time from Emilia Romagna.
Keywords: Crustacea, Decapoda, Brachyura, Pliocene, Emilia Romagna, N Italy.
]]> Resumen
Se reportan cuatro especies de cangrejos braquiuros del Plioceno (Piacenziano) de las Montañas Apeninas, todas recolectadas en drenajes naturales alrededor de Castellarano y Monticelli di Quattro Castella, Reggio Emilia (Emilia Romagna, N Italia). Estas especies se conocen en el registro fósil de Italia, y son asignadas a Calappa granulata (Linnaeus, 1758) (Calappidae De Haan, 1833); Chlinocephalus demissifrons (Ristori, 1886) (Euryplacidae Stimpson, 1871); Goneplax rhomboides (Linnaeus, 1758) (Goneplacidae MacLeay, 1838); y Monodaeus bortolottii (Delle Cave, 1988) (Xanthidae MacLeay, 1838). Chlinocephalus demissifrons y Monodaeus bortolottii se reportan por vez primera para Emilia Romagna, N de Italia.
Palabras clave: Crustacea, Decapoda, Brachyura, Plioceno, Emilia Romagna, Italia.
1. Introduction and geological setting
Many authors have reported decapod crustaceans from the Pliocene and Pleistocene of Emilia Romagna (Ristori, 1886; Vinassa de Regny, 1897; Tettoni, 1923; Beschin and Santi, 1997; Garassino and Fornaciari, 2000; Garassino and De Angeli, 2004; Garassino et al., 2010; De Angeli et al., 2011). The specimens have been discovered in two different localities along the Apennine Reggiano Mountains around Castellarano and Monticelli di Quattro Castella, both located in Reggio Emilia Province (Emilia Romagna, N Italy) (Figure 1).
1.1. Castellarano
Eight specimens (MSNM i27756, i27757, i27761, i27762, i27763, i27764, i27765, i27766) were collected from Castellarano (44°31'00"N, 10°44'00"E), around 20 km southern of Reggio Emilia, where light blue clays crop out in an old quarry today abandoned. No detailed stratigraphic data are available for this deposit, which belongs to the "Argille di Lugagnano" Formation (Cita et al., 2006). The lower circalittoral to epibathyal mollusk assemblage, however, is generally comparable, with some differences, to an assemblage from the near Campore outcrop (Parma, Emilia Romagna) from the Pliocene (Piacenzian) (Ceregato et al., 2007). Bertolaso and Garilli (2009: 15) suggested "an Early to Middle Pliocene age could be prudentially assigned" to the Castellarano deposit. Based on these data, we ascribe the specimens to the Piacenzian (Pliocene). An unidentified in situ paguroid and Retropluma craverii (Crema, 1895) (Retroplumidae Gill, 1894) were recently reported from Castellarano by Pasini and Garassino (2010) and De Angeli et al. (2011).
]]> 1.2. Monticelli di Quattro CastellaMonticelli di Quattro Castella (44°38'00"N, 10°28'00"E), western Reggio Emilia, is located among the last hills of the Apennine Mountains toward the Pianura Padana (Lombardy). Five specimens (MSNM i27758, i277 59, i27760, i27766, i27767) and six loose dactyli, were collected from some small blocks of blue clays detached by natural erosion along a gully, located in front of an old, disused quarry (Cava Moja). The scarce macrofauna assemblage includes mainly small mollusks and some irregular echinoids (Schizaster sp.) preserved as internal moulds. Lacking detailed stratigraphic data for this locality, the only geo-paleontological reports useful are those related to the study of the nearby Cava Moja (Marasti and Raffi, 1977; Monegatti and Raffi, 2001) where the malacofauna assemblage includes also deep-water forms referred to as Piacenzian (lowermost Pliocene) assemblage (Bertolaso pers. comm., 2009). Pasini and Garassino (2012) reported from this locality two cirolanid isopods (Palaega sp. and Palaega steatopigia Pasini and Garassino, 2012), referred to the Piacenzian and suspected to indicate deep-water deposits. We ascribe the specimens to the Piacenzian (Pliocene), based on these data.
2. Material
The specimens are preserved as three-dimensional moulds in small blocks of blue clays, in dorsal or ventral views, partially articulated, slightly compressed, and mostly lacking the dorsal epicuticle. Some incomplete chelipeds and one carapace were collected loose in the naturally washed sediments. The specimens have been assigned to Calappa granulata (Linnaeus, 1758) (Calappidae De Haan, 1833) (1 specimen), Chlinocephalus demissifrons Ristori, 1886 (Euryplacidae Stimpson, 1871) (1 specimen); Goneplax rhomboides (Linnaeus, 1758) (4 specimen) (Goneplacidae MacLeay, 1838); Monodaeus bortolottii Delle Cave, 1988 (Xanthidae MacLeay, 1838) (5 specimen). Moreover one specimen is assigned to an indeterminate Goneplacinae (Goneplacidae MacLeay, 1838). Some loose dactyli not reported here could be assigned to unidentified carcinids. The specimens are deposited in the Palaeontological Collections of the Museo di Storia Naturale di Milano (MSNM). The systematic arrangement used in this paper follows the recent classifications proposed by Castro (2007), Ng et al. (2008) and Schweitzer et al. (2010).
2.1. Abbreviations
lcxp: length of carapace; ld: length of dactylus; wcxp: width of carapace.
3. Systematic Paleontology
Section Eubrachyura de Saint Laurent, 1980 ]]> Superfamily Calappoidea De Haan, 1833
Family Calappidae De Haan, 1833
Genus Calappa Weber, 1795
Type species: Cancer granulatus Linnaeus, 1758, subsequent designation by Latreille (1810).
Included fossil species: see Schweitzer et al. (2010).
Calappa granulata (Linnaeus, 1758)
Cancer granulatus Linnaeus, 1758: 627.
Cancer granulatus – Linnaeus, 1767: 533.
Calappa granulata – Garassino and De Angeli, 2004: 38, fig. 4 (1-3). — Garassino et al., 2004: 264, fig. 7 a-c. — De Angeli and Garassino, 2006: 40. — Ng et al., 2008: 48. — De Angeli et al., 2009: 176, 177, 195, 196, fig. 8a, b. — Schweitzer et al., 2010: 82. — Garassino et al., 2012: 51. — Garassino and Pasini, 2013: 330, 331. — Baldanza et al., 2013: 344.
]]> Stratigraphic range: Piacenzian (Pliocene - Recent).Locality: Monticelli di Quattro Castella (Reggio Emilia).
Material and measurements: one complete right dactylus in lateral view (MSNM i27767 - ld: 10 mm).
Discussion: Although the specimen appears incomplete, the general shape of the curved dactylus, with a typical strong posterior outer lateral tooth, shows morphological affinities with Calappa granulata (Linnaeus, 1758). This species was reported from the Pliocene of Candelo, Masserano, Cossato, and S. Pietro (Piedmont) (Garassino et al., 2004; Garassino and Pasini, 2013); Arda River (Emilia Romagna) (Garassino and De Angeli, 2004); Orciano, Presciano, and Terre Rosse (Tuscany) (Ristori, 1891a; De Angeli et al., 2009); and Altavilla (Sicily) (Gemmellaro, 1914); from the Early Pleistocene of Poggio i Sodi (Tuscany) (Baldanza et al., 2013). This is the first record for this species from Reggio Emilia Province.
Superamily Goneplacoidea MacLeay, 1838
Family Euryplacidae Stimpson, 1871
Genus Chlinocephalus Ristori, 1886
Type species: Chlinocephalus demissifrons Ristori, 1886, by monotypy.
Included fossil species: Chlinocephalus demissifrons Ristori, 1886.
]]> Remarks: Schweitzer et al. (2010) erroneously reported two species as belonging to this genus, C. demissifrons Ristori, 1886, and C. subovalis (Ristori, 1886). Indeed Ristori (1886) described the new genus Chlinocephalus, with only the type species C. demissifrons; any species, named subovalis is reported in the text. The genus is therefore monotypic, as correctly reported by Karasawa and Kato (2003).Chlinocephalus demissifrons Ristori, 1886
Figure 2
Chlinocephalus demissifrons Ristori, 1886: 101, Pl. 2 (figs. 5-6).
Chlinocephalus demissifrons – Glaessner, 1929: 113. — Glaessner, 1969: R517. — Karasawa and Kato, 2003: 139, Tab. 5. — Garassino et al., 2004: 275-278, figs. 15, 16. — De Angeli and Garassino, 2006: 64. — De Angeli et al., 2009: 195. — Schweitzer et al., 2010: 133. — Garassino et al., 2012: 52.
Stratigraphic range: Piacenzian (Pliocene).
Locality: Castellarano (Reggio Emilia).
Material and measurements: one complete carapace in dorsal view, partially compressed, preserving the right cheliped and incomplete walking legs (MSNM i27756 - lcxp: 23 mm; wcxp: 25 mm; ld = 9 mm).
Discussion: The specimen reflects the main morphological characters of Chlinocephalus demissifrons Ristori, 1886, as follows: suboval, convex, smooth carapace, slightly wider than long; well-developed fronto-orbital region, wide straight front, with a weak median incision and deep wide orbits; two sharp pointed spines on anterolateral margins; convex posterolateral margin; wide, straight posterior margin; dorsal region not distinct, with two transverse ridges, marking a dorsal depression on the carapace among them; suboval, wide cardiac region; well-developed P1 with elongate merus; subcilindrical carpus with a strong spine on inner distal margin; robust chelae with elongate curved dactylus. Chlinocephalus demissifrons was already reported from the Pliocene of Fornaci (Savona, Liguria, NW Italy) (Ristori, 1886) and Biella (Piedmont, NW Italy) (Garassino et al., 2004). Moreover, Garassino et al. (2004) pointed out that the specimen assigned by Ristori (1891a) to Titanocarcinus sculptus Ristori, 1891, from the Pliocene of Mucigliani (Siena, Tuscany) could be a juvenile stage of C. demissifrons. The specimen represents the first report for the species from Emilia Romagna and the second report along the southern paleo-Adriatic Gulf.
]]>Family Goneplacidae MacLeay, 1838
Subfamily Goneplacinae MacLeay, 1838
Genus Goneplax Leach, 1814
Type species: Ocypoda bispinosa Lamarck, 1801, by original designation.
Fossil species: G. gulderi Bachmayer, 1953; G. rhomboides (Linnaeus, 1758).
Remarks: Karasawa and Kato (2003, Table 6) and Schweitzer et al. (2010) provided check lists of the fossil species of Goneplax. At present, G. craverii Crema, 1895, is assigned to Retropluma (De Angeli et al., 2011). Müller (1993) pointed out that G. formosa Ristori, 1886, and G. meneghinii Ristori, 1886, from the Pliocene of Rapolano (Siena, Tuscany) must be considered as junior synonyms of G. rhomboides (Linnaeus, 1758). Moreover, Müller (1993), comparing the main characters of the holotypes of G. sacci Crema, 1895 (G. saccoi [sic] in Karasawa and Kato, 2003, and Schweitzer et al., 2010) and G. gulderi Bachmayer, 1953, from the Miocene of Austria, pointed out the similarity of the fronto-orbital margin between the two species, different from G. rhomboides for the presence of two well-marked transverse ridges on the dorsal surface of carapace and for the front as wide as the orbits. The lost of the holotype of G. sacci questions the systematic validity of this species that should be considered as nomen dubium (for discussion, see Garassino et al., 2013). Finally, G. arenicola has been assigned to a new genus inside the Goneplacinae (Garassino et al., 2013). So based on the above-mentioned observations, we recognize only two valid fossil species of Goneplax (Garassino et al., 2013).
]]> Cancer rhomboides Linnaeus, 1758: 626.Goneplax rhomboides (Linnaeus, 1758)
Figure 3
Goneplax angulata Leach, 1814: 430.
Goneplax impressa Desmarest, 1817: 504, 505.
Goneplax romboides [sic] – Glaessner, 1929: 199. — Vía Boada, 1933: 226.
Goneplax cf. rhomboides – Gemmellaro, 1914: 90, Pl. 1 (fig. 26).
Goneplax angulata – Heller, 1863: 103. — Nobre, 1936: 57, Pl. 21 (fig. 40). — Bouvier, 1940: 278, fig. 176, Pl. 9 (fig. 2). — Zariquiey Álvarez, 1946: 162, Pl. 18 (figs. a, b).
Gonoplax [sic] impressa – Desmarest, 1822: 102–104, Pl. 8 (figs. 13, 14).
Goneplax impressa – Garassino et al., 2012: 46.
Gonoplax [sic] bispinosa – Ristori, 1891b: 20.
Stratigraphic range: Piacenzian (Pliocene - Recent).
]]> Locality: Castellarano (Reggio Emilia).Material and measurements: one incomplete specimen in dorsal view (MSNM i27762 – lcxp: 19 mm; wcxp: 25 mm); two incomplete compressed moulds (MSNM i27764, i27765) in ventral and dorsal view and one three-dimensional isolate carapace (MSNM i27766 – lcxp: 14 mm; wcxp: 20 mm).
Discussion: Although poorly preserved and partially compressed, the specimens show a subrectangular carapace with smooth dorsal surface, wider than long, widest at junction between antero- and posterolateral margins; well-developed orbits; outer orbital angle with acute spine; small acute anterolateral spine; dorsal regions not distinct; elongate chelipeds and dactyli; very elongate, flat ambulatory legs. These characters and the shape and arrangement of the pleonal sternites (where observables) allow ascribing the specimens to the fossil and extant G. rhomboides. This species has been already reported from the Pliocene-Pleistocene of Piedmont, Emilia Romagna, Tuscany, and Lazio (Ristori, 1891b; Garassino and De Angeli, 2004; Garassino et al., 2004; De Angeli et al., 2009; Garassino et al., 2012; Baldanza et al., 2013).
Genus and sp. indet.
Figure 4
Stratigraphic range: Piacenzian (Pliocene).
Locality: Castellarano (Reggio Emilia).
Material and measurements: one incomplete carapace three-dimensionally preserved in dorsal view (MSNM i27763 – lcxp: 20 mm; wcxp: 20 mm).
Description: Subquadrate carapace, as wide as long, strongly inflated in transverse section; dorsal surface of the carapace with small pits, without clear indications of regions; wide, straight front, protruding downwards, not marked by median notch; wide orbits, with supraorbital margin conspicuously sinuous; outer orbital tooth strongly projecting outwardly; straight anterolateral margin so that the fronto-orbital margin as wide as than maximum width of carapace at junction of antero- posterolateral margins; one reduced anterolateral tooth on each side of the carapace; posterolateral margins slightly rounded; straight posterior margin.
]]> Remarks: The family Goneplacidae, as reported by Castro (2007), has the following diagnostic characters: carapace transversely rectangular, subquadrate, or trapezoidal; front narrow to wide, typically lamellar, straight; dorsal surface smooth; varying number of anterolateral spines posterior to outer orbital angle (sometimes none but typically one or two); orbits moderately to conspicuously wide, long; fissure in supraorbital margin absent; dorsal surface of carapace typically smooth or with slight horizontal ridges, moderately convex, without clear indication of regions. Most of these characters are present in the specimen, here assigned to Goneplacidae. The subquadrate carapace, as wide as long, excludes the belonging of the specimen to the extant and fossil Carcinoplax H. Milne Edwards, 1852, Eutricoplax Castro, 2007, Menoplax Castro, 2007, Thyraplax Castro, 2007, Goneplax Leach, 1814, Goneplacoides Castro, 2007, Hadroplax Castro, 2007, Neogoneplax Castro, 2007, Paragoneplax Castro, 2007, Singhaplax Serène and Soh, 1976, Microgoneplax Castro, 2007, Ommatocarcinus White, 1852, Exopheticus Castro, 2007, and Neommatocarcinus Takeda and Miyake, 1969, having transversely rectangular carapace, slightly or much wider than long or subcircular carapace. The subquadrate carapace, moderately convex, without clear indication of regions is a character shared with Pycnoplax Castro, 2007, and Notonyx A. Milne-Edwards, 1873. The single anterolateral tooth on each side of carapace, however, in the specimen excludes its inclusion in Pycnoplax with two anterolateral teeth, or to Notonyx without anterolateral teeth. The Goneplacidae, as reported by Karasawa and Kato (2003) and Schweitzer et al. (2010), includes eight fossil genera, Amydrocarcinus Schweitzer, Feldmann, González-Barba and Vega, 2002; Carcinoplax H. Milne Edwards, 1852; Goneplax Leach, 1814; Icriocarcinus Bishop, 1988; Kowaicarcinus Feldmann, Schweitzer, Maxwell and Kelley, 2008; Magyarcarcinus Schweitzer and Karasawa, 2004; Ommatocarcinus White, 1852; Psopheticus Wood-Mason, 1892. Carcinoplax, Goneplax, and Ommatocarcinus are extant genera from which the specimen is excluded. Amydrocarcinus was described from the Eocene Tepetate Fm. (Mexico) (Schweitzer et al., 2002; Schweitzer and Karasawa, 2004), Icriocarcinus from the Late Cretaceous (late Campanian or early Maastrichtian) of San Diego County (California, USA) (Bishop, 1988), Kowaicarcinus from the Pliocene of New Zealand (Feldmann et al., 2008), Magyarcarcinus from the Late Eocene of Hungary (Schweitzer and Karasawa, 2004). We exclude the specimen from these genera not only for geological age and paleogeographic implications, but also for its ovoid carapace, small and squared orbits, and the dorsal surface of the carapace typically smooth of Amydrocarcinus; the pentagonal carapace, anterolateral margins with three acute spines, and the dorsal surface of the carapace strongly ridged of Icriocarcinus; the hexagonal carapace, small and ovoid orbits, and tri-lobed anterolateral margin of Kowaicarcinus; the circular carapace, small and ovoid orbits, the smooth dorsal surface of the carapace of Magyarcarcinus. Instead, Psopheticus from the Oligocene of Taiwan (Hu and Tao, 1996; Karasawa and Kato, 2003), and from the Late Pliocene of Japan (Karasawa, 1997) is the only genera having more close characters with the specimen. However the poorly preservation of the carapace and the lack of other comparative material, can not make possible to assign the specimen to this genus.Superfamily Xanthoidea MacLeay, 1838
Family Xanthidae MacLeay, 1838
Subfamily Euxanthinae Alcock, 1898
Genus Monodaeus Guinot, 1967
Type species: Xanto couchii Couch, 1851, by original designation.
Included fossil species: Monodaeus bortolottii Delle Cave, 1988.
Remarks: Schweitzer et al. (2010) reported two fossil species belonging to Monodaeus. We point out that Monodaeus couchii (Couch, 1851), however, has never been reported as a fossil, being only an extant species.
]]> Monodaeus bortolottii Delle Cave, 1988
Figure 5
Monodaeus bortolottii Delle Cave, 1988: 123-126, Pl. 1 (figs. 1, 2), Pl. 2 (figs. 1-5).
Monodaeus bortolottii – De Angeli & Garassino, 2006: 74. — De Angeli et al., 2009: 185, 195, fig. 16. — Schweitzer et al., 2010: 126. — Garassino et al., 2012: 52. — Baldanza et al., 2013: 347, 348, fig. 13.
Stratigraphic range: Piacenzian (Pliocene - late Gelasian-early Calabrian (Pleistocene).
Locality: Castellarano and Monticelli di Quattro Castella (Reggio Emilia).
Material and measurements: two incomplete carapaces (MSNM i27757, i27761 – Castellarano); three carapaces preserved in dorsal view as internal moulds including part of the chelipeds (MSNM i27758 – lcxp: 20 mm; wcxp: 25 mm.; ld: 15 mm; i27759, i27760) from Monticelli di Quattro Castella.
Discussion: Although preserved as internal moulds, the specimens show morphological affinities with Monodaeus bortolottii Delle Cave, 1988. The suboctagonal carapace is slightly convex, wider than long; straight front margin, with a weak median incision; short, convex anterolateral margin; long, convergent posterolateral margin; posterior margin straight medially and convex on margins with a granular ridge; dorsal region well marked by grooves, with wide, raised epigastric lobes; suboval protogastric regions well marked; subpentagonal mesogastric regions with narrow, elongate anterior process between protogastric regions; cardiac region well marked by branchiocardiac grooves; small hepatic regions poorly marked; wide branchial regions well marked; subtrapezioidal palm of chelipeds, wider anteriorly with a strong pointed dactylus slightly curved downwards. A rim of pointed denticles runs dorsally on the dactyli, decreasing in size towards the smooth tip. Monodaeus bortolottii has been previously reported only from the Pliocene of Volterra (Delle Cave, 1988) and Grosseto (De Angeli et al., 2009) in Tuscany. Finally, according to Baldanza et al. (2013) this species is also reported from the Early Pleistocene of Poggio i Sodi (Siena, Tuscany). This is the first record of this species from Emilia Romagna Region (N Italy).
Acknowledgments
]]> We wish to thank L. Bertolaso (Società Reggiana di Scienze Naturali, Reggio Emilia, Italy), who supplied us with useful information on geology and faunal assemblage of the localities; J.-M. Pacaud (Département Histoire de la Terre, Muséum national d'Histoire naturelle, Paris) for useful suggestions about the Roles of Zoological Code; P. Castro (Biological Sciences Department, California State Polytechnic University, Pomona, USA) for useful suggestions about the systematics of Goneplacidae, and F. J. Vega (Instituto de Geología, Universidad Nacional Autónoma de México, Ciudad Universitaria, Coyoacán, México) for careful review and criticism.
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