A New Bathyal Shark Fauna from the Pleistocene Sediments of Fiumefreddo (Sicily, Italy)

A new bathyal shark fauna from the Pleistocene sediments of Fiumefreddo (Sicily, Italy)

Stefano MARSILI Dipartimento di Scienze della Terra, Università di Pisa, Via S. Maria, 53, I-56126 Pisa (Italy) [email protected] and marsili76@@gmail.com

Marsili S. 2007. — A new bathyal shark fauna from the Pleistocene sediments of Fiumefreddo (Sicily, Italy). Geodiversitas 29 (2) : 229-247.

ABSTRACT A new shark fauna is described from the lower-middle Pleistocene marine sediments outcropping near the Fiumefreddo village, Southern Italy. Th e fossil assemblage mostly consists of teeth belonging to the bathydemersal and bathypelagic elasmobranch species Chlamydoselachus anguineus, Apristurus sp., Galeus cf. melastomus, Etmopterus sp., Centroscymnus cf. crepidater, Scymnodon cf. ringens, Centrophorus cf. granulosus and C. cf. squamosus, commonly recorded along the extant outer continental shelf and upper slope. From a bathymetrical KEY WORDS point of view, the present vertical distribution of the taxa collected allows an Elasmobranchii, estimate of a depth between 500 and 1000 m. Th e bathyal character of the sharks, bathyal, fauna provides new evidence of a highly diversiﬁ ed and heterogeneous deep-sea teeth, Mediterranean Plio-Pleistocene marine fauna, in response to the climatic and paleobiogeography, hydrographical changes. Finally, the Fiumefreddo fauna provides new relevant Pleistocene, Sicily, data for the understanding of the processes involved in the evolution of the Italy. extant Mediterranean selachian fauna.

RÉSUMÉ Une nouvelle faune de requins bathyaux des sédiments du Pléistocène de Fiume- freddo (Sicile, Italie). Une nouvelle faune de sélaciens bathyaux est décrite des sédiments marins du Pléistocène inférieur-moyen de Fiumefreddo, Italie méridionale. L’association fossile comprend en majorité des dents attribuées à des espèces d’élasmobranches bathydémersaux et bathypélagiques Chlamydoselachus anguineus, Apristurus sp.,

GEODIVERSITAS • 2007 • 29 (2) © Publications Scientiﬁ ques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 229 Marsili S.

Galeus cf. melastomus, Etmopterus sp., Centroscymnus cf. crepidater, Scymnodon cf. ringens, Centrophorus cf. granulosus et C. cf. squamosus, largement répartis actuellement sur le plateau continental externe et sur le talus supérieur. En ce qui concerne la bathymétrie, la distribution verticale actuelle des taxa a permis MOTS CLÉS d’estimer une profondeur d’environ 500 à 1000 m. Le caractère bathyal de la Elasmobranchii, faune fournit de nouvelles preuves de la mise en place d’une faune marine pro- requins, bathyal, fonde méditerranéenne au Plio-Pléistocène fortement diversiﬁ ée et hétérogène, dents, en réponse aux changements climatiques et hydrographiques. En conclusion, la paléobiogéographie, faune de Fiumefreddo fournit de nouveaux éléments importants pour la com- Pléistocène, Sicile, paraison des processus évolutifs principaux impliqués dans la mise en place de Italie. la faune actuelle méditerranéenne de sélaciens.

INTRODUCTION Th e elasmobranches represent an ecologically relevant group in the marine environments, because Th e Mediterranean Sea represents a semi-enclosed of their prominent trophodynamic. However, the basin, characterized by a very impoverished bathyal studies of this predatory group have never reached fauna. Th is is mostly due to the restricted con- a relevant role for the understanding of the main nection with the Atlantic Ocean, as well as to the Plio-Pleistocene paleoenvironmental changes of the peculiar hydrological conditions. Recently it has Mediterranean basin. Th e relatively poor knowl- been demonstrated that no more than 8.6% of the edge of the Mediterranean Plio-Pleistocene shark total Mediterranean fauna is restricted to depths fauna (see e.g., Landini 1977; Cigala Fulgosi & below 200 m (Bouchet & Taviani 1992). In the Mori 1979; Cigala Fulgosi 1984, 1988a; Cappetta last decades, studies on fossil marine vertebrate and & Nolf 1991; Bellocchio et al. 1991; Cappetta & invertebrate groups have allowed to infer the main Cavallo 2006; Carnevale et al. 2006; Marsili 2007) Plio-Pleistocene climatic and oceanographic changes only provides some data about the diversity of some and geological events that deeply characterized the epipelagic and upper mesopelagic groups. As far as evolutionary history of the Mediterranean Sea. In Mediterranean fossil deep-water elasmobranches particular, many eff orts have been made to defi ne are concerned, although bathyal taxa have been the great diversity, disparity, and heterogeneity that occasionally recorded in Pliocene sediments (see characterized the Plio-Pleistocene invertebrate and e.g., Lawley 1876; De Stefano 1911; Landini 1977; vertebrate fauna of the Mediterranean (see e.g., Cappetta 1987; Cappetta & Nolf 1991; Cappetta & Benson 1972; Landini & Menesini 1978, 1986; Cavallo 2006), well developed deep-sea assemblages Gaetani & Saccá 1984; Van Harten 1984; Raffi are rare. Cigala Fulgosi (1986, 1996), for example, 1985; Sprovieri 1985; Nolf & Cappetta 1989; Bar- described a deep-sea Mediterranean shark teeth fauna rier et al. 1989; Landini & Sorbini 1993, 2005a, b; from the lower Pliocene sediments, of Sant’Andrea Di Geronimo & La Perna 1996, 1997a; Nolf et al. a Bagni (Parma district, Northern Apennines). A 1998; Monegatti & Raffi 2001; Girone & Varola stratigraphic gap in the middle-upper Pliocene and 2001; Di Geronimo et al. 2003; Girone 2003; the Pleistocene, characterizes the fossil record of Girone et al. 2006). bathyal shark species.

230 GEODIVERSITAS • 2007 • 29 (2) Pleistocene bathyal sharks from Southern Italy

Calabria Tyrrhenian Sea

Reggio Calabria Palermo Fiumefreddo Fiumefreddo Etna Sicily Catania Etna Gela Ionian Sea Giarre

0 m 150

Acireale Apenninic-Maghrebian units

500 m Ionian Sea Volcaniclastic deposits (Holocene) N Acicastello Etna volcanics (middle Pleistocene-Holocene)

Marine sediments (lower-middle Pleistocene)

Catania 10 km Alluvial coastal plain (Recent)

FIG. 1. — Geological sketch map of Fiumefreddo (Sicily, Italy); modiﬁ ed from Lanzafame et al. (1999) and Di Stefano & Branca (2002).

Here, a new fossil shark assemblage has been stud- MATERIALS AND METHODS ied from the marine sediments outcropping near the Fiumefreddo village (Catania district, Sicily). Ten fossiliferous horizons were examined. Th e sam- Th is fossil assemblage improves our understanding ples (20 dm3, routinely processed and sieved for of the relationships existing between the climatic macrofaunal studies) were collected for the study of and hydrographical changes and the development fi sh otoliths fauna (see Girone et al. 2006), and then of a diversifi ed bathyal Plio-Pleistocene fauna in the provided to the author for the study of the shark Mediterranean Sea (see e.g., Barrier et al. 1989; Di teeth. Th irty-four fossil teeth were extracted from Geronimo & La Perna 1996, 1997a; Girone et al. eight of these fossiliferous horizons (see Table 1). 2006), presenting new data for understanding the About half of the teeth collected are well preserved, evolutionary and historical processes involved in while the remaining material is fragmentary crowns the establishment of the extant Mediterranean fi sh and roots of little taxonomic interest. Th is sample fauna (see also Landini & Sorbini 2005a, b). included a number of teeth belonging to the teleostean

GEODIVERSITAS • 2007 • 29 (2) 231 Marsili S.

family Trichiuridae, from several horizons (see Table 1999; Di Stefano & Branca 2002). Th e presence of 1). Th e specimens are deposited in the Museo di the foraminiferas Hyalinea balthica and Globorotalia Paleontologia of the University of Catania (PMC). truncatulinoides excelsa suggested an age for these Morphological and tooth terminology mainly fol- marine sediments of lower-middle Pleistocene lows that of Ledoux (1970), Cappetta (1987), and (Lombardo 1980; Lanzafame et al. 1999). Recently, Herman et al. (1989, 1990, 1993). Taxonomic Di Stefano & Branca (2002), on the basis of the information were derived from papers by Garrick nannoplancton calcareous content, referred the (1956, 1959a-c), Cigala Fulgosi & Gandolfi (1983), lower part of this section to the “Small” Gephyro- Compagno (1984), Cigala Fulgosi (1988b, 1996) capsa Zone of Rio et al. (1990), and the uppermost and Adnet & Cappetta (2001). part to the Pseudoemiliana lacunose Zone of Rio et al. (1990). Th e fossil material documented in this paper GEOLOGICAL SETTING has been collected from a section located near the Fiumefreddo village (Catania district, Sicily), along Th e northern and southern sector of the Etna Vol- the northern fl ank of the Serra S. Biagio hill (Fig. 1). cano are characterized by a continuous bioclastic Th e succession is represented by about 4.5 m of calcarenites and marly clays succession (Lanzafame silty clays conformably covered by about 5 m of et al. 1999; Di Stefano & Branca 2002). In the blue marly clays (Angela Girone pers. comm.). Th e southern sector, the calcarenites and marly clays shark teeth described herein were extracted from sequence is part of a Plio-Pleistocene regressive seven silty clays horizons (Fiumefreddo Nord 2-8), succession that characterized the basement of the belonging to the “Small” Gephyrocapsa Zone, and Gela-Catania foredeep (Lanzafame et al. 1999; Di one marly clays horizon (Fiumefreddo Cava 2) re- Stefano & Branca 2002). In the north-eastern sec- ferred to the Pseudoemiliana lacunose Zone (Girone tor, such a marine succession unconformably covers et al. 2006). the basement, represented by the inner units of the Apenninic-Maghrebian Chain (Lanzafame et al. 1999; Di Stefano & Branca 2002). A complex SYSTEMATICS geodynamic evolution, that characterized the frontal area of the Apenninic-Maghrebian Chain, caused a Order HEXANCHIFORMES Buen, 1926 more consistent uplift of the Pleistocene deposits Family CHLAMYDOSELACHIDAE Garman, 1884 of the northern sector, as a consequence of a buried Genus Chlamydoselachus Garman, 1884 thrust located on the northern fl ank of the Etna Volcano (Di Stefano & Branca 2002). Chlamydoselachus anguineus Garman, 1884 Th e most complete calcarenites and marly clays (Fig. 2A-D) section crops out along the fl anks of the Serra S. Biagio hill, in the northern sector of the Etna MATERIAL EXAMINED. — One incomplete tooth and one Volcano (Lombardo 1980; Lanzafame et al. 1997, eroded root (PMC Fiumefreddo 1996. I. Pl. Sel. 1, 2).

FIG. 2. — A-D, Chlamydoselachus anguineus Garman, 1884, anterior-lateral tooth (PMC Fiumefreddo 1996. I. Pl. Sel. 1); A, lingual view; B, lateral view; C, labial view; D, basal view; E-J, Centrophorus cf. granulosus (Bloch & Schneider, 1801); E, upper anterior tooth, lingual view (PMC Fiumefreddo 1996. I. Pl. Sel. 12); F, G, upper anterior-lateral tooth (PMC Fiumefreddo 1996. I. Pl. Sel. 13); F, labial view; G, lingual view; H, I, lower anterior-lateral tooth (PMC Fiumefreddo 1996. I. Pl. Sel. 14); H, lingual view; I, labial view; J, lower anterior-lateral tooth, lingual view (PMC Fiumefreddo 1996. I. Pl. Sel. 15); K-N, Scymnodon cf. ringens Bocage & Capello, 1864; K, L, upper anterior-lateral tooth (PMC Fiumefreddo 1996. I. Pl. Sel. 10); K, labial view; L, lingual view; M, N, lower anterior- lateral tooth (PMC Fiumefreddo 1996. I. Pl. Sel. 11); M, labial view; N, lingual view; O, P, Etmopterus sp., lower anterior-lateral tooth (PMC Fiumefreddo 1996. I. Pl. Sel. 8); O, lingual view; P, labial view; Q, R, U, Centrophorus cf. squamosus (Bonnaterre, 1788); Q, R, dermal scale, ventral view; Q, PMC Fiumefreddo 1996. I. Pl. Sel. 16; R, PMC Fiumefreddo 1996. I. Pl. Sel. 17; U, dermal scale, lateral view (PMC Fiumefreddo 1996. I. Pl. Sel. 17); S, T, Centroscymnus cf. crepidater (Bocage & Capello, 1864), lower commissural tooth (PMC Fiumefreddo 1996. I. Pl. Sel. 9); S, labial view; T, lingual view. Scale bars: 3 mm.

232 GEODIVERSITAS • 2007 • 29 (2) Pleistocene bathyal sharks from Southern Italy

ABCD

EF G H I

JKLMN

OP QR

ST U

GEODIVERSITAS • 2007 • 29 (2) 233 Marsili S.

FIG. 3. — Apristurus aff. laurussoni (Saemundsson, 1922), lower anterior tooth (PMC Fiumefreddo 1996. I. Pl. Sel. 3): A, lingual view; B, labial view. Scale bar: 1 mm.

DESCRIPTION by two high, thick crests. Th e central concave to Th e incomplete tooth lacks the distal lateral cusp and slightly fl at region is characterized by a large central part of the root. Th e median and mesial lateral cusps foramen. Th e distal and mesial regions are fl at and are thin, slender and sigmoid, with a few short basal slightly sloped, with few small foramina randomly folds. Th e mesial lateral cusp is more sigmoid than scattered. Two or three small foramina are present the median one and laterally divergent. Th e labial on the inner end of the lingual crests. face of the cusps is fl at to slightly convex, while the lingual face is completely convex. Th e lateral mesial REMARKS and distal cutting edges of the cusps are smooth. Th e characters described, the very peculiar shape, Th ere is a very short intermediate cusplet between as well as its strict morphological affi nity with the the median and mesial lateral cusps. extant teeth published by Bass et al. (1975a: pl. 6) Th e root is perpendicular to the cusps. Th e labial and Herman et al. (1993: pls 1, 2) support the assign- face of the root is short with a well developed basal ment of the specimens to the Recent C. anguineus. mesial depression. Th e basal face of the root is broad, Lawley (1876: pl. 1, fi g. 1-1b) fi rst described fos- fl at, and labio-lingually elongated with two lobe-like sil teeth of Chlamydoselachus, later on assigned by extensions directed lingually, that provided a trans- Davis (1887) to the fossil taxon C. lawleyi Davis, versal interlocking with the labial mesial depressions 1887. Since no teeth of living or fossil species were of the root of the previous tooth of the same row. A known to Lawley (1876), he only provided a brief mesial depression characterized the basal face of the description of the specimens, without any taxonomic root, in which is presents a large central oval-shape assignment. Th e extant species was described only foramen. A great number of small foramina are ran- few years later by Garman (1884a, b, 1885). Th e domly scattered all over the root base. Th e lingual diagnosis proposed by Davis (1887) was based on an face of the root is divided into three distinct regions indirect comparison between the fossil tooth drawn

234 GEODIVERSITAS • 2007 • 29 (2) Pleistocene bathyal sharks from Southern Italy

by Lawley (1876) and a single extant specimen of continuous from the crown base towards the apex, C. anguineus. Davis (1887: 544) remarked that the are also present on the labial face of cusplets. fossil teeth was twice the diameter of the anterior Th e root is high, with a well pronounced torus teeth of the extant specimens. Unfortunately, the characterized by a large central lingual foramen, holotype of C. lawleyi has been lost, so that the from which a deep basal central transverse groove taxonomic validity of the morphometric characters originates. A few small lingual foramina (3-4) are used by Davis cannot be stated defi nitively. More- present on the lingual face of the root lobes. Th e over, C. lawleyi diff ers from C. anguineus because basal face of the root is fl at and labio-lingually of the absence of the intermediate cusplets and it elongated, divided by the basal central transverse may have been omitted by Lawley himself (Richter groove into two distinct lobes. A large number of & Ward 1990). Th e uncertainty of the characters small basal foramina are randomly scattered on the used to defi ne C. lawleyi and its morphological basal surface of the root lobes. Th e labial face of the affi nities with the teeth of living species suggests root is low and partially eroded, with a longitudi- that the fossil taxon might be conspecifi c with nal central depression characterized by fi ve large C. anguineus (Richter & Ward 1990). labial foramina. Th e more central labial foramina are merged in the anterior tooth.

Order CARCHARHINIFORMES REMARKS Compagno, 1973 Th e presence of few linear to sigmoid folds on the Family SCYLIORHINIDAE Gill, 1862 labial face of the narrow and elongated principal Genus Apristurus Garman, 1913 cusp, continuous from the crown base toward the apex, as well as the presence of a deep central trans- Apristurus sp. verse groove on the basal face of the root (Herman (Fig. 3) et al. 1990) justify the assignment of the studied samples to the genus Apristurus. Moreover, Apris- MATERIAL EXAMINED. — One incomplete lower anterior turus sp. shares a close tooth morphology with the tooth and one mostly incomplete lower lateral tooth (PMC Fiumefreddo 1996. I. Pl. Sel. 3, 4). teeth of the living A. laurussoni (Saemundsson, 1922) described by Herman et al. (1990: pls 1-4). DESCRIPTION However, the taxonomic problems concerning this Th e anterior teeth are characterized by a narrow genus (see Iglésias & Nakaya 2004), as well as the and elongated principal cusp and by two mesial poor extant comparative materials do not allow a and distal lateral cusplets. Th e specimens studied specifi c assignment of the specimens studied. preserve only the mesial lateral cusplets, of which the closest reaches half the height of the principal cusp. Th e lateral tooth is incomplete and presents Genus Galeus Rafi nesque, 1810 three mesial lateral cusplets of which the closest reaches almost the height of the bent principal cusp. Galeus cf. melastomus Rafi nesque, 1810 Th e lingual faces of the principal cusp and cusplets (Fig. 4) are convex, and present some folds, more evident along the basal region of the face than near the Galeus cf. melastomus – Cigala Fulgosi 1986: 135. apex. Th e labial face of the principal cusp is fl at to MATERIAL EXAMINED. — Two lower anterior-lateral teeth slightly convex in the basal region, and completely and one lower lateral tooth (PMC Fiumefreddo 1996. convex in the upper region, characterized by 6 or I. Pl. Sel. 5, 6, 7). 7 linear to sigmoid folds, much more evident than those present on the lingual face of the cusp. Th e DESCRIPTION more central labial folds run from the crown base Th e anterior tooth is characterized by a narrow towards the tip of the cusp. A few folds (1 or 3), and elongated principal cusp, and by two mesial

GEODIVERSITAS • 2007 • 29 (2) 235 Marsili S.

AB C D

FIG. 4. — Galeus melastomus Rafi nesque, 1810: A, B, lower anterior tooth (PMC Fiumefreddo 1996. I. Pl. Sel. 5); A, lingual view; B, labial view; C, D, lower anterior-lateral tooth (PMC Fiumefreddo 1996. I. Pl. Sel. 6); C, lingual view; D, labial view. Scale bars: 1 mm. and distal lateral cusplets, of which the closest to the living Galeus cf. melastomus. Th is deep-water reach half the height of the principal cusp, while shark shares a very close tooth morphology with the distal ones reach half the height of the closest Apristurus. However the presence of obvious folds cusplets. Th e lateral tooth is characterized by two on the labial face of the principal cusp that never distal and three mesial lateral cusplets, of which reach the crown apex (Herman et al. 1990), as well the closest reach almost the height of the strongly as the absence of a basal deep central transverse bent principal cusp. Th e lingual faces of the cusp groove (Cappetta 1987; Herman et al. 1990) are and cusplets are convex, and present a few folds, the main characters to separate G. cf. melastomus more evident along the basal region of the face from Apristurus sp. than near the apex. Th e labial face of the principal Fossils of the genus Galeus have been found in cusp is fl at to slightly convex, and characterized by the Burdigalian and Langhian bathyal deposits of 6 or 7 folds that run from the crown base toward France (Cappetta 1987), and the living species was half the height of the labial face of the cusp, never fi rstly recorded in the lower Pliocene deposit of the reaching the tip of the cusp. By contrast, a few Northern Apennines (Cigala Fulgosi 1986). Th e continuous folds (1 or 3) are present on the labial presence of G. cf. melastomus in the Pleistocene face of the cusplets. sediments has been recently confi rmed also in the Th e root is high, with a well pronounced torus, Italian section of Archi and Vallone Catrica, in the characterized by a large central lingual foramen. A Reggio Calabria district (unpublished data). few small lingual foramina (3 or 4) are present on the lingual face of the root lobes. Th e basal face of the root is fl at and labio-lingually elongated, with Order SQUALIFORMES Goodrich, 1909 a large number of small randomly scattered basal Suborder DALATIOIDEI foramina. Th e basal central deep transverse groove is sensu De Carvalho 1996 completely absent. Th e labial face of the root is low, Family DALATIIDAE Gray, 1851 with a longitudinal central depression characterized Genus Etmopterus Rafi nesque, 1810 by 5 or 6 large labial foramen. Th e more central labial foramina are merged in the anterior tooth. Etmopterus sp. (Fig. 2O, P) REMARKS According to Herman et al. (1990), the characters MATERIAL EXAMINED. — One complete lower anterior- described justify the assignment of the specimens lateral tooth (PMC Fiumefreddo 1996. I. Pl. Sel. 8).

236 GEODIVERSITAS • 2007 • 29 (2) Pleistocene bathyal sharks from Southern Italy

DESCRIPTION eastern Atlantic species E. princeps Collet, 1904 Th e tooth is labio-lingually compressed. Th e cusp is or E. pusillus Lowe, 1839 does not allow a specifi c strongly inclined and narrow, with the tip slightly assignment of the material studied. turned upward. Th e mesial cutting edge is wavy Th e presence of Etmopterus sp. in the Pleistocene and completely smooth. Th e distal cutting edge Italian sediments has been recently confi rmed also is short and convex, separated by a deep notch in the Archi section, in the Reggio Calabria district from a distal smooth enamel blade, convex in the (unpublished data). outline. Th e apron is rectangular and wide, and it completely covers the upper central and distal region of the labial face of the root. Th e uvula is Genus Centroscymnus Bocage & Capello, 1864 rectangular and wide, and it completely covers the upper central and mesial region of the lingual face Centroscymnus cf. crepidater of the root. (Bocage & Capello, 1864) Th e root is rectangular and incomplete. Th e lin- (Fig. 2S, T) gual face of the root is fl at, with a deep strong distal depression, resulting from interlocking of the teeth. Centroscymnus cf. crepidater – Cigala Fulgosi 1986: 134. A thick longitudinal ridge is present just below the MATERIAL EXAMINED. — One complete lower commis- basal margin of the uvula. Th ree lingual foramina sural tooth (PMC Fiumefreddo 1996. I. Pl. Sel. 9). are present along the uvula-root junction, of which the mesial one is much larger than the central and DESCRIPTION distal foramina. A small lingual central foramen Th e tooth is labio-lingually compressed, with a characterized also the lower region of the lingual short and inclined cusp. Th e mesial cutting edge is face of the root. Th e labial face of the root is fl at, sigmoid and completely smooth. Th e distal cutting with a strong mesial depression, resulting from edge is perpendicular, short and completely smooth, interlocking of the teeth. Two labial foramina are separated by a deep notch from a long distal enamel present between the basal margin of the apron and blade. Th e apron is rectangular, projected down- the root, while two other labial foramina are present ward, reaching half the height of the labial face of between the mesial margin of the apron and the the root. Th e uvula is completely absent. distal margin of the mesial depression of the root. Th e root is rectangular, and mesio-distally elon- A small incomplete central labial foramen is also gated, characterized by an irregular and oblique present in the lower region of the labial face. distal basal margin. Th e lingual face of the root is fl at, characterized by a longitudinal ridge near the REMARKS crown-root junction, on which a large mesial and Th e presence of uvula and apron that cover the whole central foramen is present. Th e lingual axial groove upper region of the root (Ledoux 1970; Herman et is absent. Th e labial face of the root is fl at, char- al. 1989) supports the assignment of the specimens acterized by a hardly mesial depression, resulting to the genus Etmopterus. Th e lower tooth collected from interlocking of the teeth. Th ree large labial in the Fiumefreddo section diff ers from the Recent foramina, vertically stacked, are present along the E. spinax (Linnaeus, 1758) because of a narrower and mesial margin of the apron. Few small labial foramina slightly more erected cusp, with a tip turned upward, are all around the distal margin of the apron and and for a lower number of labial foramina along the crown-root junction. A shallow and short axial the apron-root junction (see Herman et al. 1989). groove characterized the lower region of the root. Th e Miocene Etmopterus sp. reported by Ledoux (1972) diff ers from the Fiumefreddo specimens in REMARKS having a shorter cusp and a much higher root. Th e Th e absence of a lingual axial groove on the lin- scant knowledge about the dental variation of the gual face of the root (Herman et al. 1989) and Mediterranean specie E. spinax, as well as of the the strong morphological affi nity with the teeth

GEODIVERSITAS • 2007 • 29 (2) 237 Marsili S.

of C. crepidater published by Ledoux (1970) and root is fl at, and forms an angle with the cusp. A Herman et al. (1989) support the assignment of the very small lingual central foramen is present along studied material to this bathydemersal shark species. the root-crown junction, while a narrow and deep Because of the general morphology of the root, the lingual axial groove characterizes the basal region specimens studied can be considered a commis- of the lingual face. Th e labial face of the root is sural tooth (see e.g., Ledoux 1970: fi g. 17Ad). A characterized by a deep central depression, in which less erected cusp in the lower teeth and the absence a narrow and deep labial axial groove is present, of a lingual axial groove on the lingual face of the and by a mesial and distal depression, in which tooth distinguish C. crepidater from Scymnodon few foramina are randomly scattered. ringens Bocage & Capello, 1864 (Ledoux 1970; Th e lower anterior-lateral tooth is labio-lingually Herman et al. 1989; Adnet & Cappetta 2001). compressed, with a triangular erected cusp distally Centroscymnus crepidater diff ers from the Mediter- inclined. Th e mesial and distal cutting edges are com- ranean specie C. coelolepis Bocage & Capello, 1864 pletely smooth. Th e mesial cutting edge is straight to and from C. cryptacantus (Bocage & Capello, 1864) slightly sigmoid. Th e distal cutting edge is straight, in having a shorter root and apron (Ledoux 1970; and separated by a deep notch from a distal enamel Herman et al. 1989). Th e same characters might be blade. Th e uvula is absent. Th e apron is rectangular, used also to separate C. crepidater from the species narrow and not so much elongated. C. owstoni Garman, 1906 and C. plunketi (Waite, Th e root is rectangular. Th e lingual face of the 1900) (Garrick 1959b, c), but more detailed and root is fl at, characterized by low longitudinal ridge exhaustive studies are necessary. along the crow-root junction, on which a mesial and central lingual foramen is present. A shallow incomplete lingual axial groove is present on the Genus Scymnodon Bocage & Capello, 1864 basal region of the lingual face of the root. Th e labial face of the root is fl at, with a well developed Scymnodon cf. ringens Bocage & Capello, 1864 mesial depression, resulting from interlocking of (Fig. 2K-N) the teeth. Several small labial foramina, vertically arranged, are present along the mesial and distal Scymnus lichia – De Stefano 1911: 398, pl. 10, fi gs margins of the apron, of which the upper are larger 27, 28. than the lower ones. Scymnodon cf. ringens – Cigala Fulgosi 1986: 135. REMARKS Scymnodon ringens – Cigala Fulgosi 1996: 314, pl. 5, fi g. 9, pl. 6, fi g. 6. Scymnodon ringens shares a close tooth similarity with the bathyal shark of the genus Zameus (sensu MATERIAL EXAMINED. — One upper anterior-lateral tooth Cigala Fulgosi 1996), Centroscymnus and Scym- and one lower anterior-lateral tooth (PMC Fiumefreddo nodalatias Garrick, 1956. An upper anterior-lateral 1996. I. Pl. Sel. 10, 11). tooth with a bilobate root (Garrick 1956; Herman et al. 1989; Cigala Fulgosi 1996) separated Scym- DESCRIPTION nodalatias from S. ringens. Centroscymnus diff ers Th e upper anterior-lateral tooth is characterized from Scymnodon in having upper teeth with one by a conical narrow, sigmoid and elongated cusp. or two large foramina on the lingual face of the Th e lateral cutting edges are smoothed and incom- root (Herman et al. 1989), and a bilobate root (see plete. Th e lingual face of the cusp is convex. Th e C. colelolepis in Ledoux 1970; Herman et al. 1989). uvula is absent. Th e labial face of the crown is fl at, Finally, S. ringens diff ers from Zameus squamulosus with a deep basal depression bordered by a thick (Günther, 1877) in having a quadrangular root mesial and distal vertical ridge, characterized by a in the upper anterior-lateral teeth and a shorter few costules. root and apron in the lower teeth (Herman et al. Th e root is quadrangular. Th e lingual face of the 1993).

238 GEODIVERSITAS • 2007 • 29 (2) Pleistocene bathyal sharks from Southern Italy

TABLE 1. — Systematic list of sharks from Fiumefreddo (Sicily, Italy) with relative total amount of specimens per sample. Fiumefreddo Fiumefreddo 2 Nord Fiumefreddo 3 Nord Fiumefreddo 4 Nord Fiumefreddo 5 Nord Fiumefreddo 6 Nord Fiumefreddo 7 Nord Fiumefreddo 8 Nord Fiumefreddo Cava 2 Chlamydoselachidae Chlamydoselachus anguineus 11 Scyliorhinidae Apristurus sp. 1 1 Galeus cf. melastomus 12 Dalatiidae Etmopterus sp. 1 Centroscymnus cf. crepidater 1 Scymnodon cf. ringes 11 Centrophoridae Centrophorus cf. granulosus 31 Centrophorus cf. squamosus 12 Fragmentary shark teeth 1 2 5 4 2 2 Teeth of teleostean family Trichiuridae 1 1 41 2

Th e upper lateral-anterior tooth is the only re- DESCRIPTION corded in the middle Pleistocene sediments of the Th e upper anterior tooth is higher than wide, char- Fiumefreddo section, and in particular it has been acterized by a more or less symmetrical cusp. Th e collected from the “Fiumefreddo Cava 2” horizon mesial lateral cutting edge is convex and smooth, (Table 1), located in the upper part of the section. while the distal one is concave and smooth. Th e Th is tooth also represents the fi rst middle Pleistocene upper anterior-lateral tooth is almost as high as record of the bathyal shark S. ringens. wide. Th e mesial lateral cutting edge is concave and smooth, while the distal one is slightly concave and smooth. Laterally to the cusp, a smoothed mesial Family CENTROPHORIDAE Bleeker, 1859 and distal enamel blades are present. Th e upper Genus Centrophorus Müller & Henle, 1837 teeth are characterized by a narrow and elongated apron, that reaches the basal margin of the labial Centrophorus cf. granulosus face of the root. A short smoothed uvula is also (Bloch & Schneider, 1801) present. (Fig. 2E-J) Th e lower anterior-lateral teeth exhibit a very distally inclined cusp. Th e mesial lateral cutting edge Acanthias major Lawley, 1876: 40, pl. 1, fi g. 19a, b. is slightly convex and poorly serrated. Th e distal Acanthias vulgaris – De Stefano 1910: 594, pl. 17, fi gs cutting edge is short and straight, separated by a 23, 24. deep notch from a distal enamel blade. Th e apron is narrow and elongated, stopping just before the Centrophorus granulosus – Ledoux 1972: 145, fi g. 5. — Landini 1977: 119, fi g. 4. basal margin of the labial face of the root. In one specimen the basal margin of the apron is branched. Centrophorus cf. granulosus – Cigala Fulgosi 1986: 134. A short smoothed uvula is present. Centrophorus aff . granulosus – Cappetta & Nolf 1991: Th e root is quadrangular in the upper anterior 52, pl. 2, fi gs 1-4. — Cappetta & Cavallo 2006: 36, tooth, and becomes sub-rectangular in the upper pl. 1, fi g. 2. anterior-lateral and in the lower anterior-lateral teeth. Th e lingual face of the root is characterized MATERIAL EXAMINED. — One upper anterior tooth; one upper anterior-lateral tooth; two lower anterior- by a depressed crown-root junction, below which a lateral teeth (PMC Fiumefreddo 1996. I. Pl. Sel. 12, thick longitudinal ridge divided the lingual surface 13, 14, 15). into an upper and lower region. A large lingual

GEODIVERSITAS • 2007 • 29 (2) 239 Marsili S.

foramen, located mesially beside the uvula, is present and uvula, and by the presence of larger foramina on the longitudinal ridge. A shallow infundibulum, on the labial and lingual faces of the root (Cigala that originates from the lingual foramen, is scarcely Fulgosi & Gandolfi 1983; Cigala Fulgosi 1988b; developed, because of the eroded root of the teeth. Herman et al. 1989). A deep distal depression, resulting from interlock- Th e presence of C. granulosus in the Pleistocene ing of the teeth, is present in the upper region of sediments has been recently confi rmed also in the the lingual face of the root. Few small foramina are Italian section of Grammichele, in the Catania randomly scattered on the upper distal and mesial district (unpublished data). region of the lingual face of the root. Th e labial face of the root is fl at, with a deep mesial depression, resulting from interlocking of the teeth. In Centrophorus cf. squamosus the upper anterior teeth, one large mesial and one (Bonnaterre, 1788) distal foramina are present laterally to the apron. (Fig. 2Q, R, U) Several small foramina are also present along the crown-root junction. In the lower teeth, along Centrophorus squamosus – Keyes 1984: 206, fi gs 8- the crown-root junction, one or two large labial 20. — Cigala Fulgosi 1986: 134. foramina, and several small labial foramina are MATERIAL EXAMINED. — Th ree dermal scales (PMC present in the upper mesial region of the labial face Fiumefreddo 1996. I. Pl. Sel. 16, 17, 18). of the root. Two or three large labial foramina, and several small labial foramina are also present on the DESCRIPTION upper distal region of the labial face. Th e dermal scales are characterized by a slender and elongated peduncle, that sustained a rhom- REMARKS boidal and leaf-like crown. Th e basal region of Th e presence of a large lingual foramen located the peduncle is widen, and adorned by few small mesially beside the uvula, and of an infundibu- crests. Th e crown is characterized by arched and lum that originates from the lingual foramen, thick anterior-lateral margins, and by straight and support the assignment of the specimens to the serrated posterior- lateral margins. Th e serrations genus Centrophorus. Th e species of this genus are of the posterior-lateral margins are character- characterized by a very similar tooth morphology ized by a posterior central strong main cusp and (Cigala Fulgosi 1986; Cappetta & Nolf 1991). by 3 or 4 small lateral cusplets. A thick central Th e strong morphological affi nity with the teeth longitudinal ridge is present on the ventral face of C. granulosus published by Ledoux (1970) and of the crown, and runs from the anterior region Herman et al. (1989) supports the assignment of of the crown to the tip of the posterior central the studied material to this bathydemersal shark main cusp. Two small lateral ridges run along species. An uvula without ornamentations (Ledoux the anterior-lateral margins of the crown, paral- 1970) separates C. granulosus from C. squamosus lel to the central longitudinal ridge, and stop at (Bonnaterre, 1788), while a larger apron and uvula the midpoint of the ventral face of the crown. (Cappetta & Nolf 1991) separates this shark from Th ese lateral ridges can be forked in two parallel C. lusitanicus Bocage & Capello, 1864. Th e teeth branches. No ornamentations are present on the of the genus Deania Jordan & Snyder, 1902 share ventral face of the crown. a close morphology with Centrophorus, except that in Deania the infundibulum is absent and two large REMARKS axial foramina are present (Ledoux 1970, 1972; Centrophorus squamulosus is known in the Pleis- Cappetta & Nolf 1991; Adnet & Cappetta 2001). tocene sediments of Fiumefreddo section only by Th e teeth of Somniosus rostratus (Risso, 1826) diff er the presence of rare dermal scales. from those of Centrophorus because they are higher Th e presence of a slender and elongated pedun- than wide, by the much wider and shorter apron cle sustaining a rhomboidal and leaf-like crown,

240 GEODIVERSITAS • 2007 • 29 (2) Pleistocene bathyal sharks from Southern Italy

TABLE 2. — Present status and paleoecological preferences of the shark taxa from Fiumefreddo (Sicily, Italy).

Taxa Present status Environment Depth range (m) Chlamydoselachus Extra-Mediterranean Bathydemersal Outer shelf-upper 0-1570 anguineus slope Apristurus sp. Extra-Mediterranean Bathydemersal Upper slope 560-1462 Galeus cf. melastomus Mediterranean Bathydemersal Outer shelf-upper 0-1873 slope Etmopterus sp. Mediterranean Bathydemersal Outer shelf-upper 0-2490 slope Centroscymnus cf. Extra-Mediterranean Bathydemersal Upper slope 230-1500 crepidater Scymnodon cf. ringes Extra-Mediterranean Bathypelagic Slope 200-1600 Centrophorus cf. Mediterranean Bathydemersal Outer shelf-upper 100-1200 granulosus slope Centrophorus cf. Extra-Mediterranean Bathypelagic Slope 145-2400 squamosus and of a crown characterized by a posterior-lateral recorded for the fi rst time in the middle Eocene margins with three or four lateral cusplets and a deposits of Landes, in the South-West of France posterior main cusp (Garrick 1959a; Hulley 1971; (Adnet 2006). Finally, the only shark tooth collected Bass et al. 1975b; Compagno 1984), supports the in the fossiliferous horizon “Fiumefreddo Cava 2” assignment of the specimens to the Recent leaf-scale (Table 1) represents the fi rst middle Pleistocene gulper shark C. squamosus. All the other species of record of Scymnodon cf. ringens. the genus Centrophorus diff er from C. squamosus by Th e assemblage is characterized by temperate to the absence of a slender peduncle and a diff erent subtropical bathydemersal and bathypelagic shark physiognomy of the crown (Compagno 1984). A species, that commonly inhabit the deep waters of rounded to ovoid or elongated crown with two or the outer continental shelves and/or upper slope three posterior main cusps distinguishes Centroscym- (Compagno 1984) (Table 2). Th e taxa identifi ed nus, Scymnodalatias and Scymnodon ringens from show the high bathyal shark diversity of the sec- C. squamosus (Bigelow & Schroeder 1957; Garrick tion, in contrast with the present Mediterranean 1956, 1959b, c; Compagno 1984). Moreover, the Sea condition, characterized by a very impoverished absence of cross-ridges on the ventral surface of the deep-water fauna (Bouchet & Taviani 1992; Sion crown excludes the possible assignment to Zameus et al. 2004). Several Atlantic taxa now absent in squamulosus. the Mediterranean are represented in this Pleis- tocene assemblage, including C. anguineus, S. cf. ringens, Centroscymnus cf. crepidater, Centrophorus DISCUSSION cf. squamosus and Apristurus sp. Th e genus Apris- turus included a lot of species with a more or less Th e Fiumefreddo shark fauna provides new knowl- restricted extra-Mediterranean geographic distri- edge of the Mediterranean Plio-Pleistocene bathyal bution (see Compagno 1984). In particular, A. elasmobranch distribution, disparity and diversity. laurussoni, characterized by a tooth morphology Th is fossil shark fauna represents the fi rst deep-sea near to Fiumefreddo specimens, is a North Atlantic tooth assemblage recorded in the lower Pleistocene species, characterized by a discontinuous distribu- sediments of the Mediterranean basin. In particular, tion along the Massachusetts and Delaware coasts, the tooth assigned to Chlamydoselachus anguineus in the Gulf of Mexico, along the south of Ireland represents the fi rst fossil record of this species. and in the Canary and Madeira islands (Compagno Moreover, the teeth assigned to Apristurus sp. fi ll 1984; Iglésias & Nakaya 2004). S. ringens is the in part the fossil record of the genus Apristurus, only extra-Mediterranean species of Fiumefreddo

GEODIVERSITAS • 2007 • 29 (2) 241 Marsili S.

section distributed only in the eastern Atlantic Ocean, Cigala Fulgosi (1996) related the disappearance from Scotland to Senegalese coasts (Compagno of many of these Atlantic bathyal taxa, just at the 1984). By contrast, C. anguineus, C. squamosus and end of the Zanclean, to a rapid normalization of C. crepidater are characterized by wider geographical the Mediterranean water circulation (estuarine distribution, recorded both in the Atlantic, Pacifi c type model circulation, deep homothermy, nu- and Indian oceans (Bass et al. 1975a, b; Nakaya & trients depletion), that led to a strong impover- Bass 1978; Compagno 1984). Etmopterus sp., Cen- ishment of the deep-water Mediterranean fauna. trophorus cf. granulosus and Galeus cf. melastomus Th e Fiumefreddo assemblage disagrees with a instead represent some of the rare bathyal sharks rapid disappearance of bathyal sharks from the characteristic of the present deep Mediterranean Mediterranean, suggesting a continuity in deep- fauna (Sion et al. 2004). water shark diversity and disparity until the lower Concerning the paleobathymetry (see Compagno Pleistocene. Moreover, the occurrence of several 1984), all the shark species collected along the bathyal shark teeth (see also Marsili & Tabanelli in Fiumefreddo horizons occur today at depths below press), as well as of other invertebrate groups (see 150-200 m. Apristurus laurussoni is characterized by e.g., Monegatti & Pantoli 1987; Tabanelli 1993; a bathymetric range of 560 m to 1462 m. C. angui- Vaiani & Venezia 1999; Monni & Tabanelli 2006; neus, C. crepidater and C. granulosus are characterized Monni 2006) from the middle-upper Pliocene of by a deeper bathymetric limit that ranges in depth Romagna, confi rms the persistence of a more At- from 1000 to 1200 m. S. ringens may reach at least lantic deep-water fauna also during these periods about 1600 m depth. C. squamosus is a rare shark in the Mediterranean. above 1000 m depth in the eastern Atlantic, and, Th e strong oceanization and biodiversity of the likewise Etmopterus sp. and G. melastomus, has a Pleistocene bathyal shark fauna from Fiumefreddo lower bathymetric limit placed at around 2000 m. agrees with a more upper Pliocene-lower Pleistocene Today, in the Mediterranean deep waters, Etmop- general Mediterranean faunistic trend, character- terus spinax, C. granulosus and G. melastomus have ized by a large and progressive entrance of both been recently recorded at depths that range from Atlantic deep-water invertebrate and vertebrate about 600 to about 1500 m, and, in the eastern marine group in the basin (see e.g., Gaetani & Ionian Sea, E. spinax has been captured at about Saccá 1984; Barrier et al. 1989; Di Geronimo & 2200 m depth (Sion et al. 2004). Th erefore, the La Perna 1996, 1997a, b; Di Geronimo et al. 1997, Fiumefreddo shark fauna supports a depositional 2003; Rosso & Di Geronimo 1998; Corselli 2001; environment with a depth between about 500 and Girone & Varola 2001; Girone 2003; Girone et 1000 m. Th is bathymetric range is in agreement al. 2006). Th e Plio-Pleistocene global climatic with the large number of Trichiurid teeth (Table 1), cool trend and the subsequent lowering of the as well as with the previous study on the otoliths median annual sea surface temperature (Dowsett fi sh fauna from the same section of Fiumefreddo & Poore 1990; Dowsett et al. 1994; Herbert & (Girone et al. 2006). Schuff ert 1998), permitted the entrance of bo- Most of the Atlantic bathyal taxa collected in the real and/or arctic cold stenothermic taxa, such as studied section appeared in the Mediterranean Sea the ostracod Cytheropteron testudo (Bonaduce & during the early Pliocene (Cigala Fulgosi 1986, Sprovieri 1985), or the molluscs Arctica islandica 1996; Cappetta & Nolf 1991). Th eir occurrence (Raffi 1985; Malatesta & Zarlenga 1986) and was favoured by the peculiar deep-water hydro- Homalopoma emulum (Di Geronimo & La Perna graphical conditions that during the lower Pliocene 1997b). A similar trend can be observed also for characterized this basin, due to the establishment some teleostean fi shes such as Nansenia groenlan- of an estuarine type water circulation, as well as dica, Bathylagus euryops, Coryphaenoides rupestris or to the presence of a lower sill in the Gibraltar Aphanopus carbo (Girone et al. 2006). Moreover, area (see e.g., Benson 1975; Th unnel et al. 1987; a deeper connection between the Atlantic Ocean McKenzie et al. 1990; Iaccarino et al. 1999). and the Mediterranean Sea, characterized by the

242 GEODIVERSITAS • 2007 • 29 (2) Pleistocene bathyal sharks from Southern Italy

absence of a real threshold at Gibraltar, favoured are thanked for their thorough reviews and helpful the persistence of these cold stenothermic taxa in suggestions. I extend special thanks to Prof. Italo this basin (Barrier et al. 1989; Di Geronimo & La Di Geronimo, and Prof. Antonietta Rosso (both Perna 1996, 1997a). Dipartimento di Scienze della Terra, Università di None of the Atlantic bathydemersal and bathy- Catania) for providing the samples from the Fiume- pelagic shark species collected in the studied section freddo, Grammichele, Archi and Vallone Catrica are characterized by an exclusive boreal or arctic sections, and to Dr. Angela Girone (Dipartimento geographic distribution, as well as a stenothermic di Scienze della Terra, Università di Bari) for provid- character. However, their presence in the Fiume- ing the samples from the Fiumefreddo section, as freddo section confi rms that diff erent hydrographic well as for her useful stratigraphical and geological conditions of the Mediterranean deep waters existed suggestions. Th e present study was granted by the during the lower Pleistocene, as well as a major “Scuola di Dottorato del Dipartimento di Scienze connection with the near Atlantic Ocean (Barrier et della Terra, University of Pisa”. al. 1989; Di Geronimo & La Perna 1996, 1997a). Moreover, such an organized elasmobranch fauna supports the establishment of a basin system char- REFERENCES acterized by a decreasing temperature, a good deep nutrient supply and a better deep-water circulation, similar to the near eastern Atlantic, that favoured ADNET S. 2006. — Nouvelles faunes de sélaciens (Elas- mobranchii, Neoselachii) de l’Éocène des Landes the development of a rich deep ecosystem in the (Sud-Ouest, France). Implication dans les connais- Mediterranean, supported by a well structured sances des communautés d’eaux profondes. Palaeo trophic system, where sharks played an important Ichthyologica 10: 1-128. role. Th e present Mediterranean hydrographical ADNET S. & CAPPETTA H. 2001. — A paleontologi- conditions (see e.g., Bouchet & Taviani 1992), cal and phylogenetical analysis of squaliform sharks (Chondrichthyes: Squaliformes) based on dental due to a diff erent circulation model (Th unell et al. characters. Lethaia 34: 234-248. 1987) and to a reached “threshold basin condition” BARRIER P. , D I GERONIMO I., MONTENAT C., ROUX in the Gibraltar region (Di Geronimo & La Perna M. & ZIBROWIUS H. 1989. — Présence de faunes 1996, 1997a), caused the disappearance of many bathyales atlantiques dans le Pliocène et le Pléistocène cold stenothermic taxa from the Mediterranean Sea, de Méditerranée. Bulletin de la Société géologique de France 8 (4): 787-796. representing today also the main extant physical and BASS A. J., D’AUBREY J. D. & KISTNASAMY N. 1975a. — biological barrier to the penetration in the basin of Sharks of the East Coast of Southern Africa. V. Th e many Recent deep-water Atlantic species. families Hexanchidae, Chlamydoselachidae, Pristio- phoridae and Squatinidae. Oceanographic Research Institute, Investigational Report 43: 1-50. BASS A. J., D’AUBREY J. D. & KISTNASAMY N. 1975b. — Acknowledgements Sharks of the East Coast of Southern Africa. VI. I would like to thank Prof. Walter Landini (Dipar- Th e families Oxynotidae, Squalidae, Dalatiidae and timento di Scienze della Terra, Università di Pisa) Echinorhinidae. Oceanographic Research Institute, for his suggestions and critical reading of an early Investigational Report 45: 1-103. draft of the text as well as for his useful suggestions. BELLOCCHIO G., CARBONI M. G., NAMI M. & PALLINI G. 1991. — Fauna ad ittiodontoliti del Pliocene di Many thanks go to Dr. Giorgio Carnevale (Dipar- Allerona (Terni, Umbria). Bollettino della Società dei timento di Scienze della Terra, Università di Pisa) Naturalisti in Napoli 100: 41-73. for his critical reading of an early draft of the text BENSON R. H. 1972. — Ostracods as indicators of and for his friendly support. Prof. Henry Cappetta threshold depth in the Mediterranean during the (Institut des Sciences de l’Évolution, Université de Pliocene, in STANLEY D. J. (ed.), Th e Mediterranean Sea: a Natural Sedimentation Laboratory. Dowden, Montpellier II) and Mr. Robert Purdy (Department Hutkinson & Ross, Stroudsburg: 63-73. of Paleobiology, National Museum of Natural His- BENSON M. L. 1975. — Th e origin of the psychro- tory, Smithsonian Institution, Washington D.C.) sphere as recorded in changes of deep-sea ostracode

GEODIVERSITAS • 2007 • 29 (2) 243 Marsili S.

assemblage. Lethaia 8: 63-73. Società Paleontologica Italiana 34 (3): 301-322. BIGELOW H. B. & SCHROEDER W. 1957. — A study of CIGALA FULGOSI F. & MORI D. 1979. — Osservazioni the sharks of the suborder Squaloidea. Bulletin of the tassonomiche sul genere Galeocerdo (Selachii, Car- Museum of Comparative Biology 117 (1): 1-150. charhinidae) con particolare riferimento a Galeocerdo BONADUCE G. & SPROVIERI R. 1985. — Th e appearance cuvieri (Péron & Lesueur) nel Pliocene del Mediter- of Cytheropteron testudo Sars (Crustacea: Ostracoda) raneo. Bollettino della Società Paleontologica Italiana is a Pliocene event. Evidence from a sicilian sequence 18 (1): 117-132. (Italy). Bollettino della Società Paleontologica Italiana CIGALA FULGOSI F. & G ANDOLFI D. 1983. — Re-descrip- 23 (1): 131-136. tion of the external morphology of Somniosus rostratus BOUCHET P. & T AVIANI M. 1992. — Th e Mediterranean (Risso, 1826), with special reference to its squamation deep-sea fauna: pseudopopulation of Atlantic species? and cutaneous sensory organs, and aspects of their Deep-Sea Research 39 (2): 169-184. functional morphology (Pisces, Selachii, Squalidae). CAPPETTA H. 1987. — Chondrichthyes II. Mesozoic Monitore Zoologico Italiano (N.S.) 17: 27-70. and Cenozoic Elasmobranchii, in SCHULTZE H. P. COMPAGNO L. J. V. 1984. — FAO species catalogue. (ed.), Handbook of Paleoichthyology. Gustav Fischer Vol. 4. Sharks of the World. An annotated and il- Verlag, Stuttgart; New York, 193 p. lustrated catalogue of shark species known to date. CAPPETTA H. & NOLF D. 1991. — Les sélaciens du FAO Fisheries Synopsis 125 (4): 1-655. Pliocène inférieur de Le-Puget-Sur-Argens (Sud-Est CORSELLI C. 2001. — Change and diversity: the Mediter- de la France). Palaeontographica Abteilung A 218: ranean deep corals from the Miocene to the present, 49-67. in FARANDA F. M . , G UGLIELMO L. & SPEZIE G. (eds), CAPPETTA H. & CAVALLO O. 2006. — Les sélaciens Mediterranean Ecosystem: Structures and Processes. du Pliocène de la région d’Alba (Piémont, Italie Springer-Verlag, Berlin; New York: 361-366. Nord-Ouest). Rivista Piemontese di Storia Naturale DAVIS J. W. 1887. — Note on a fossil species of Chlamy- 27: 33-76. doselachus. Proceedings of the Zoological Society of CARNEVALE G., MARSILI S., CAPUTO D. & EGISTI L. London 1887: 542-544. 2006. — Th e silky shark Carcharhinus falciformis DE CARVALHO M. R. 1996. — Higher level elasmo- (Bibron, 1841) in the Pliocene of Cava Serredi (Fine branch phylogeny, basal squaleans, and paraphyly, Basin, Italy). Neues Jahrbuch für Geologie und Paläon- in STIASSNY M. L. J., PARENTI L. R. & JOHNSON G. tologie, Abhandlungen 242 (2/3): 257-370. D. (eds), Interrelation of Fishes 3. Academic Press, CIGALA FULGOSI F. 1984. — Contributo alla conoscenza New York: 35-62. della fauna ad elasmobranchi del Pliocene del Medi- DE STEFANO G. 1910. — Osservazioni sulla ittiofauna terraneo. Segnalazione di Notorynchus (Euselachii, pliocenica di Orciano e San Quirico in Toscana. Bol- Hexanchidae) nella “Collezione Lawley”. Acta Natu- lettino della Società Geologica Italiana 28: 539-648. ralia de “L’Ateneo Parmense” 20: 85-96. DE STEFANO G. 1911. — Sui pesci pliocenici dell’imolese. CIGALA FULGOSI F. 1986. — A deep water elasmo- Bollettino della Società Geologica Italiana 29: 381- branch fauna from a lower Pliocene outcropping 402. (Northern Italy), in UYENO T., ARAI R., TANIUCHI DI GERONIMO I. & LA PERNA R. 1996. — Bathyspinula T. & MATSUURA K. (eds), Indo-Pacifi c Fish Biology: excisa (Philippi, 1844) (Bivalvia, Protobranchia): a Proceedings of the Second International Conference on witness of the Plio-Quaternary history of the deep Indo-Pacifi c Fishes. Ichthyological Society of Japan, Mediterranean benthos. Rivista Italiana di Paleonto- Tokyo: 133-139. logia e Stratigrafi a 102 (1): 105-118. CIGALA FULGOSI F. 1988a. — Additions to the Eocene DI GERONIMO I. & LA PERNA R. 1997a. — Pleistocene and Pliocene fi sh fauna of Italy. Evidence of Alopias cf. bathyal molluscan assemblages from Southern Italy. deniculata Cappetta, 1981 in the Bartonian-Priabon- Rivista Italiana di Paleontologia e Stratigrafi a 103 (3): ian of the Monte Piano Marl (Northern Apennines) 389-426. and of A. superciliosus (Lowe, 1840) in the Pliocene DI GERONIMO I. & LA PERNA R. 1997b. — Homalo- of Tuscany (Chondrichthyes, Alopiidae). Tertiary poma emulum (Seguenza) a bathyal cold stenothermic Research 10 (2): 93-99. gastropod in the Mediterranean Pleistocene. Geobios CIGALA FULGOSI F. 1988b. — Addition to the Pliocene 30 (3): 215-223. fi sh fauna of Italy. Evidence of Somniosus rostratus DI GERONIMO I., D’ATRI A., LA PERNA R., ROSSO A., (Risso, 1826) from the foothills of the Northern SANFILIPPO R. & VIOLANTI D. 1997. — Th e Pleis- Apennines (Parma Province, Italy) (Chondrichthyes, tocene bathyal section of Archi (Southern Italy). Bol- Squalidae). Tertiary Research 10 (2): 101-106. lettino della Società Paleontologica Italiana 36 (1-2): CIGALA FULGOSI F. 1996. — Rare oceanic deep water 189-212. squaloid sharks from the Lower Pliocene of the North- DI GERONIMO I., DI GERONIMO R., ROSSO A., GIRONE ern Apennines (Parma Province, Italy). Bollettino della A. & LA PERNA R. 2003. — Autochthonous and

244 GEODIVERSITAS • 2007 • 29 (2) Pleistocene bathyal sharks from Southern Italy

allochthonous assemblage from Lower Pleistocene tocene fi sh otoliths from the Mediterranean basin: a sediments (Palione River, Sicily). Bollettino della Società synthesis. Geobios 39 (5): 651-671. Paleontologica Italiana 42 (1-2): 133-138. HERBERT T. D. & SCHUFFERT J. D. 1998. — Alkenone DI STEFANO A. & BRANCA S. 2002. — Long-term uplift unsuturation estimates of late Miocene through late rate of the Etna Volcano basement (southern Italy) Pliocene sea-surface temperature at Site 958. Proceed- based on biochronological data from Pleistocene ings of the ODP, Scientifi c Results 159T: 17-21. sediments. Terra Nova 14: 61-68. HERMAN J., HOVESTADT-EULER M. & HOVESTADT D. C. DOWSETT H. J. & POORE R. Z. 1990. — A new planktic 1989. — Contribution to the study of the comparative foraminifer transfer function for estimating Pliocene- morphology of teeth and other relevant ichthyodoru- Holocene paleoceanographic conditions in the North lites in living superspecifi c taxa of Chondrichthyan Atlantic. Marine Micropaleontology 16: 1-23. fi shes. Part A: Selachii. No. 3: Order: Squaliformes DOWSETT H. J., THOMPSON R. S., BARRON J. A., CRONIN – Families: Echinorhinidae, Oxynotidae and Squali- T. M., FLEMING F., ISHNAM S., POORE R. Z., WILLARD dae. Bulletin de l’Institut royal des Sciences naturelles D. & HOLTZ J. R. T. 1994. — Joint investigations de Belgique, Biologie 59: 101-158. of the Middle Pliocene Climate I: PRISM paleoen- HERMAN J., HOVESTADT-EULER M. & HOVESTADT vironmental reconstructions. Global and Planetary D. C. 1990. — Contribution to the study of the Science 9: 169-195. comparative morphology of teeth and other relevant GAETANI M. & SACCÀ D. 1984. — Brachiopodi ba- ichthyodorulites in living superspecifi c taxa of Chon- tiali nel Pliocene e Pleistocene di Sicilia e Calabria. drichthyan fi shes. Part A: Selachii. No. 2b: Order: Rivista Italiana di Paleontologia e Stratigrafi a 90 (3): Carcharhiniformes – Families: Scyliorhinidae. Bulletin 407-458. de l’Institut royal des Sciences naturelles de Belgique, GARMAN S. 1884a. — “Th e oldest living type of Verte- Biologie 60: 181-230. brata”, Chlamydoselachus. Science 3: 345. HERMAN J., HOVESTADT-EULER M. & HOVESTADT GARMAN S. 1884b. — Th e oldest living type of verte- D. C. 1993. — Contribution to the study of the brates. Science 3: 484. comparative morphology of teeth and other rel- GARMAN S. 1885. — Chlamydoselachus anguineus Garm. evant ichthyodorulites in living superaspecifi c taxa A living species of cladodont shark. Bulletin of the of Chondrichthyan fi shes. Addendum to Part A, No. Museum of Comparative Zoology 12 (1): 1-35. 1b: Hexanchiformes – Family: Chlamydoselachidae; GARRICK J. A. F. 1956. — Studies on New Zealand No. 5: Order : Heterodontiformes – Family: Hetero- Elasmobranchii – Part V. Scymnodalatias n.g. Based dontidae; No. 6: Order: Lamniformes – Families: on Scymnodon scherwooi Archey, 1921 (Selachii). Cetorhinidae, Megachasmidae; Addendum 1 to no. Transaction of the Royal Society of New Zealand 83: 3: Order: Squaliformes; Addendum 1 to no. 4: Order 555-571. Orectolobiformes; General Glossary; Summary Part GARRICK J. A. F. 1959a. — Studies on New Zealand A. Bulletin de l’Institut royal des Sciences naturelles de Elasmobranchii. Part VII. Th e identity of Centrophorus Belgique, Biologie 63: 185-256. from New Zealand. Transactions of the Royal Society HULLEY P. A. 1971. — Centrophorus squamosus (Bon- of New Zealand 86: 127-141. naterre) (Chondrichthyes, Squalidae) in the Eastern GARRICK J. A. F. 1959b. — Studies on New Zealand South Atlantic. Annals of the South African Museum Elasmobranchii – Part VIII. Two northern hemi- 57 (11): 265-270. sphere species of Centroscymnus in New Zealand IACCARINO S., CASTRADORI D., CITA M. B., DI STEFANO waters. Transaction of the Royal Society of New Zea- E., GABOARDI S., MCKENZIE J. A., SPEZZAFERRI S. land 87: 75-89. & SPROVIERI R. 1999. — Th e Miocene-Pliocene GARRICK J. A. F. 1959c. — Studies on New Zealand boundary and the signifi cance of the earliest fl ooding Elasmobranchii – Part IX. Scymnodon plunketi (Waite, in the Mediterranean. Memorie della Società Geologica 1910), an abundant deep-water shark of New Zea- Italiana 54: 109-131. land waters. Transaction of the Royal Society of New IGLÉSIAS S. P. & NAKAYA K. 2004. — Apristurus atlanticus Zealand 87: 271-282. (Koefoe, 1927), a junior synonym of A. laurussoni GIRONE A. 2003. — Th e Pleistocene bathyal teleostean (Saemudsson, 1922) (Chondrichthyes: Carcharhini- fauna of Archi (Southern Italy): paleoecological and formes: Scyliorinidae). Cybium 28 (3): 217-223. paleobiogeographic implication. Rivista Italiana di KEYES I. W. 1984. — New records of fossil elasmobranch Paleontologia e Stratigrafi a 109 (1): 99-110. genera Magascyliorhinus, Centrophorus, and Dalatias GIRONE A. & VAROLA A. 2001. — Fish otoliths from the (Order Selachii) in New Zealand. New Zealand Journal Middle Pleistocene deposits of Montalbano Jonico of Geology and Geophysics 27: 203-216. (Southern Italy). Bollettino dell Società Paleontologica LANDINI W. 1977. — Revisione degli “Ittiodontoliti Italiana 40: 431-443. pliocenici” della collezione Lawley. Palaeontografi a GIRONE A., NOLF D. & CAPPETTA H. 2006. — Pleis- Italica 70: 92-134.

GEODIVERSITAS • 2007 • 29 (2) 245 Marsili S.

LANDINI W. & MENESINI E. 1978. — L’ittiofauna plio- MCKENZIE J. A., SPROVIERI R. & CHANNEL J. E. T. pleistocenica della sezione della Vrica (Crotone, Ca- 1990. — Th e terminal Messinian fl ood and earliest labria). Bollettino della Società Paleontologica Italiana Pliocene paleoceanography in the Mediterranean: 17 (2): 143-175. results from ODP Leg 127, Site 652, Tyrrhenian Sea. LANDINI W. & M ENESINI E. 1986. — L’ittiofauna Memorie della Società Geologica Italiana 44: 81-91. pliocenica della sez. di Stuni ed i suoi rapporti con MONEGATTI P. & P ANTOLI D. 1987. — First evidence of l’ittiofauna plio-pleistocenica della Vrica (Crotone, Pleurotomariidae (Gastropoda, Prosobranchia) from Calabria). Bollettino della Società Paleontologica Ital- the Mediterranean Pliocene. Bollettino della Società iana 25 (1): 41-63. Paleontologica Italiana 25 (3): 313-316. LANDINI W. & SORBINI L. 1993. — Biogeographic and MONEGATTI P. & RAFFI S. 2001. — Taxonomic diver- paleoclimatic relationships of the Middle Pliocene sity and stratigraphic distribution of Mediterranean ichthyofauna of the Samoggia Torrent (Bologna, Italy). Pliocene bivalves. Palaeogeography, Palaeoclimatology, Proceedings of the First RCANS Congress 12: 83-89. Palaeoecology 165: 171-193. LANDINI W. & S ORBINI C. 2005a. — Evolutionary MONNI S. 2006. — Analisi delle associazioni a molluschi trends in the Plio-Pleistocene ichthyofauna of the in una sezione pliocenica nei pressi di Brisighella (Ra- Mediterranean Basin: nature, timing and magnitude venna). Quaderni di Studi e Notizie di Storia Naturale of the extinction events. Quaternary International della Romagna 22: 1-38. 131: 101-107. MONNI S. & TABANELLI C. 2006. — Eumelia alicei LANDINI W. & SORBINI C. 2005b. — Evolutionary dy- (Dautzemberg & Fisher, 1886) ospite atlantico nel namics in the fi sh fauna of the Mediterranean Basin Pliocene romagnolo. Quaderni di Studi e Notizie di during the Plio-Pleistocene. Quaternary International Storia Naturale della Romagna 19: 1-5. 140-141: 64-89. NAKAYA K. & BASS A. J. 1978. — Th e frill shark Chlamy- LANZAFAME G., LEONARDI A., NERI M. & RUST D. doselachus anguineus in New Zealand seas. New Zea- 1997. — Late overthrust of the Appenine-Maghre- land Journal of Marine and Freshwater Research 12 (4): bian Chain at the NE periphery of Mt. Etna, Italy. 397-398. Comptes Rendus de l’Académie de Sciences de Paris sér. NOLF D. & CAPPETTA H. 1989. — Otolithes de poissons 2a, 234: 325-332. pliocènes du Sud-Est de la France. Bulletin de l’Institut LANZAFAME G., LEONARDI A. & NERI M. 1999. — Retro- royal des Sciences naturelles de Belgique, Science de la ricoprimento medio-pleistocenico di Argille Scagliose Te r re 58: 209-271. a Serra San Biagio (Sicilia orientale): evidenze strati- NOLF D., MAÑÉ R. & LOPEZ A. 1998. — Otolithes grafi che e tettoniche. Rendiconti dell’Accademia dei de poissons du Pliocène inférieur de Papiol, près de Lincei, Scienze Fisiche e Naturali 9: 63-80. Barcelone. Palaeovertebrata 27 (1-2): 1-17. LAWLEY R. 1876. — Nuovi Studi Sopra i Pesci ed altri Ver- RAFFI S. 1985. — Th e signifi cance of marine boreal tebrati Fossili delle Colline Toscane. Firenze, 122 p. mollusks in the Early Pleistocene faunas of the Medi- LEDOUX J.-C. 1970. — Les dents des squalidés de la terranean Sea. Palaeogeography, Palaeoclimatology, Méditerranée occidentale et de l’Atlantique Nord- Palaeoecology 52: 267-289. Ouest africain. Vie et Milieu, série A: Biologie marine, RICHTER M. & WARD D. J. 1990. — Fish remains (extrait) 21 (2A): 309-362. from the Santana Formations (Late Cretacous) of LEDOUX J.-C. 1972. — Les Squalidae (Euselachii) mio- James Ross Island, Antarctica. Antarctic Science cènes des environs d’Avignon (Vaucluse). Documents 2 (1): 67-76. des Laboratoires de Géologie de la Faculté des Sciences RIO D., RAFFI I. & VILLA G. 1990. — Pliocene-Pleis- de Lyon 52: 133-175. tocene calcareous nannofossil distribution patterns in LOMBARDO G. 1980. — Stratigrafi a dei depositi della the Western Mediterranean. Proceedings of the ODP, Sicilia nord-orientale. Bollettino dell’Accademia Gioenia Science Research 107: 513-533. di Scienze Naturali di Catania 14: 85-113. ROSSO A. & DI GERONIMO I. 1998. — Deep-sea Pleis- MALATESTA A. & ZARLENGA F. 1986. — Northern guests tocene Bryozoa of Southern Italy. Geobios 30 (3): in the Pleistocene Mediterranean Sea. Geologica Ro- 303-317. mana 25: 91-154. SION L., BOZZANO A., D’ONGHIA G., CAPEZZUTO F. MARSILI S. 2007. — Revision of the teeth of the genus & PANZA M. 2004. — Chondrichthyes species in Carcharhinus (Elasmobranchii; Carcharhinidae) from the deep waters of the Mediterranean Sea. Scientia the Pliocene of Tuscany, Italy. Rivista Italiana di Pale- Marina 68 (suppl. 3): 153-162. ontologia e Stratigrafi a 113 (1): 79-95. SPROVIERI R. 1985. — Paleotemperature changes and MARSILI S. & TABANELLI C. in press. — Bathyal sharks speciation among benthic Foraminifera in the Mediter- from the middle Pliocene of the Romagna Apennines ranean Pliocene. Bollettino della Società Paleontologica (Italy). Neues Jahrbuch für Geologie und Paläontologie, Italiana 24: 13-21. Abhandlungen. TABANELLI C. 1993. — Osservazioni e ipotesi sulle

246 GEODIVERSITAS • 2007 • 29 (2) Pleistocene bathyal sharks from Southern Italy

malacofaune plioceniche della Romagna. Quaderni tocenica del Lamone (Appennino romagnolo): as- di Studi e Notizie di Storia Naturale della Romagna sociazione a foraminiferi ed evoluzione paleoambi- 2: 1-20. entale. Bollettino della Società Paleontologica Italiana THUNELL R. C., WILLIAMS D. F. & HOWELL M. 1987. — 38 (1): 39-57. Atlantic-Mediterranean water exchange during the late VAN HARTEN D. 1984. — A model of estuarine circu- Neogene. Paleoceanography 2 (6): 661-678. lation in the Pliocene Mediterranean based on new VAIANI S. C. & VENEZIA P. 1999. — La sezione pleis- ostracod evidence. Nature 312: 359-361.

Submitted on 20 November 2006; accepted on 2 March 2007.

GEODIVERSITAS • 2007 • 29 (2) 247