Cainozoic Research, 3-23, March 2009 6(1-2), pp.

A small fossil fish fauna, rich in Chlamydoselachus teeth, from

the Late Pliocene of Tuscany (Siena, central Italy)

4 Franco Cigala+Fulgosi ¹*, Simone Casati Alex Orlandini³ & Davide Persico ²,

'Dipartimento di Scienze della Terra, V.le G.P. Usberti, 157/A, Campus Universilario, 43100. Parma, Italia.

Address for correspondence: Str. Martinellan. 292, 43100 Parma, Italia. E-mail: [email protected] (* correspondingauthor).

2 via Celestino Bianchin. 18, 50134 Firenze, Italia.

3 via Lombroso Cesare n. 15, 42100 Reggio Emilia, Italia.

4 Dipartimento di Scienze della Terra, V. le G.P. Usberti, 157/A, Campus Universilario, 43100, Parma, Italia.

Received 17 January 2007; revised version accepted 30 July 2008

A small elasmobranch teeth and teleost otolithassemblage from the Piacenzian-earliest Gelasian ofTuscany (Castelnuovo Berardenga

Scalo, Siena province, Italy) is described. The exceptional abundance of teeth belonging to Chlamydoselachus lawleyi Davis, 1887 has enabled us to better defineand confirmthe validity ofthis doubtful fossil taxon. In agreement with Davis (1887) and Pfeil (1983) this

the shark C. 1884 least its species appears to differ from living frilled anguineus Garman, at by larger size.

Palaeoecological inferences based onthe ecology ofC. anguineus together with deep water sharks such as Centrophorus granulosus

(Bloch & Schneider, 1801) and the teleost otolith assemblage mainly characterised by macrourids and myctophids enable us to configure,

in the Piacenzian earliest Gelasian of subbasin of the Siena-Radicofani to Tuscany (Siena Basin) an upper bathyal slope palaeo-

environment. The occurrenceofChlamydoselachus in the Piacenzian-earliestGelasian ofthe Mediterranean palaeo-area demonstrates the

of oceanic environment persistence an not as profusely developed as during the early Zanclean (i.e. lower part of MPL 2 foraminiferal biozone; see Cigala Fulgosi, 1986, 1996).

Viene descrittaunafauna a denti di elasmobranchi ed otoliti di teleostei rinvenuta nel Piacenziano-Gelasianoantico della Toscana (Cas-

telnuovoBerardengaScalo, provincia di Siena, Italia centrale).L’eccezionale abbondanza di denti di Chlamydoselachus lawleyi Davis,

1887 ha di definire incerto fossile e di confermarne la validità. permesso meglio questo taxon In accordo con Davis (1887) e Pfeil (1983)

dal dal collare C. 1884 almeno la sensibilmente questa specie differisce vivente squalo anguineus Garman, per taglia piu grande.

Induzioni basate di Cen- paleoecologiche sull’ecologia Chlamydoselachus anguineusGarman, 1884e di squali di acque profonde come

trophorus granulosus (Bloch & Schneider, 1801) e sulla associazione fossile ad otoliti di teleostei caratterizzata eminentemente da

macruridi e mictofidipermette di configurate nel Piacenziano Gelasiano antico della Toscana (sub-bacino di Siena del Bacino Siena-

Radicofani) unpaleoambiente epibatiale di scarpata. Chlamydoselachus indica inoltrela persistenza nel Piacenziano Gelasiano basale

della mediterranea di “oceanizzazione” che tuttavia paleoarea una non appare cos) intensacome quella dello Zancleano antico (i.e. parte

inferiore della biozona a foraminiferi MPL 2; Cigala Fulgosi, 1986, 1996).

KEY WORDS: fauna, Elasmobranchs, Teleosts, : Chlamydoselachus Teeth, Otoliths, Late Pliocene, Tuscany, Central Italy.

Introduction no longer possible to execute additionalcollecting.

Since the beginning of paleo-ichthyology the Pliocene

A small elasmobranch tooth and fish otolith fauna shark fauna of has been discussed unusu- Tuscany by numerous rich in of is ally teeth Chlamydoselachus described. The scholars (e.g. Lawley, 1875,Principi, 1920). Robert Law-

with few hundred surface- teeth, together a otoliths, were ley in particular played an important role, both because of

in S. Casati in collected, recent years, by an open quarry the pioneering and systematic use of the methodology,

near the village ofCastelnuovo Berardenga Scalo, Asciano based on a comparison of fossils with living beings, and

The (Siena Province) (Figure 1). quarry exploiting the because of the intensity with which he focussed his atten-

clays belongs to “Laterizi Arbia SPA” and is close to the tion on this fauna. An original biographical research about

NE side ofthe processing plant. A first attempt to process this author has recently been published by Manganelli etal.

the surface sediment in search of micro-ichtyodontoliths (2006). Many finds reported by Lawley, some of which

unsuccessful. Due the accumulation of was to recent trans- were exceptional and belonged to his original collection,

ported material on top of the outcropping sediment this have unfortunately been lost.

has been and the time it is operation not repeated at present Itwas Lawley (1876, pi. 1, fig. 1) who, while ignoring their -4-

thesis), using the occurrence of the planktonic foraminifer

GloborotaliaaemilianaColalongo & Sartoni, 1967, attrib-

uted the marls ofthe Castelnuovo Berardenga Scalo quarry

to the “Middle Pliocene”.

Nannofossil biostratigraphy

Material and methods — A semiquantitative calcareous

nannofossil analysis has been done by Davide Persico. Six

recovered samples were along ten topographic meters of

the in the stratification slope outcrop. The on the surface of

the small fossiliferous area appears indistinct. Nannofossil

preservation is good and the sediment sampling is charac-

1. location of the with terized sand and Figure Map showing outcrop Chlamy- by clay components together with the bio-

Castelnuovo Siena doselachus, at Berardenga Scalo, prov- stratigraphic markers. ince, Italy.

nature, published the first pictures of a fossil tooth of

Chlamydoselachus. Garman (1884) described this unusual shark a few years later. The nine teeth Lawley collected,

“two perfectly complete, and seven more or less mutilated”

(“due perfettamente completi, e sette piu o meno mutilati”,

Lawley, 1876, p. 87), came from Orciano Pisano (Pisa province, Italy), a village at approximately 80 km from

Castelnuovo Berardenga Scalo. It is also important to bear in mind that at that time the geographical references relat- ing to the locationsof the finds were not as precise as they are today and that very often other people collected and obtainedthese materials. According to shark teeth amateurs and collectors, the Pliocene-Pleistocene landscape around

Orciano is now developed, intensively cultivated and/or urbanized. It is therefore uncertain whether we can inter-

finds prete Lawley’s original stratigraphically (Landini,

1977). They could be generally indicated as Zanclean-

Piacenzian(“Lower-Middle Pliocene”, according to Land- ini et al., 2005 and Bianucci & Landini, 2005). Other than

Lawley’s pictures, the Pliocene species Chlamydoselachus lawleyi until now represented by fragments from other ,

Mediterraneanareas only (Pfeil, 1983), can be tested using this new abundantand mate- exceptionally well-preserved Figure 2. Chronostratigraphic scale and geomagnetic polarity rial. time scale from Lourens et al. (1996, 2004); calcareous nan-

nofossil biostratigraphy from Rio et al. (1990).

Geological setting

The adopted chronostratigraphic scheme (Figure 2) has

The small where the teeth and otoliths col- been the scale of outcrop were composed by magnetostratigraphic lected is composed of muddy and silty marls comprised Lourens et al. (2004), correlated with the biostratigraphy of withinthe subbasin ofSiena(SB), which, together with the Rio et al. (1990). Calcareous nannofossils were examined

Radicofani subbasin (RB), forms the Siena-Radicofani Ba- using standard light microscope techniques, under crossed sin (SRB) (Bonini, 2002, fig. 2). This basin extends in a polarizers, transmitted light, and phase contrast light at roughly NNW-SSE direction, and the northern part (par- 1250x magnification. The species considered have been ticularly the SB), is boundedby ridges mostly uplifted by distributed in five levels ofabundance (Table 1).

W to WSW dipping thrust faults. The sedimentary succes-

sion and ofthe basin the — stratigraphy was reported by same Results The recognized calcareous nannofossil assem-

divided the succession into four main author who angular blage characterizes a biostratigraphic interval within the unconformities-bounded stratigraphic units (Bonini, 2002, Discoaster tamalis- and Discoaster pentaradiatus-zones fig. 4). Previously, Augugliaro (1999-2000, unpublished (Figure 2). -5 -

1111 o jo o o o o Systematic Palaeontology 0) |Q) Q) 0) Q) Q) Sample J2-S2.S5.J2.S2.S2. CD_ CD_ CD_ CD_

3 3 3 3 3 3 C C C C C C Given that the fish faunaaccompanying Chlamydoselachus CastelnuovoO OCastelnuovo OCastelnuovo CastelnuovoO CastelnuovoO CastelnuovoO <<<<<< o o o o o o does not present particular taxonomic novelties, and that

® 03 03 TO 03 CD 0 SpeciesP taxa co co : i» o) the are chrono- B.S.6 wB.S.5 B.S.4 B.S.3 B.S.2co B.S.1 represented nearly always Neogene b> oi gj ro and/or that species lineages canbe related to living species,

R Amaurolithus delicatus we shall only describe their essential characters. The syn-

which reduced to dis- FF F R onymies, are a minimum, are partly Braarudosphaera bigelowi cussed in more or less recent works {e.g. Landini, 1977; C C C Cc C Calcidiscus leptoporus Compagno, 1984, 2001;Pfeil, 1983;Ward& Galea Bona-

R R F Coccolithus pelagicuspelagicus via, 2001; Antunes & Cappetta, 2002). The dental termi-

is the one used authors R R R RR R nology adopted largely by (e.g. Discoaster asymmetricus Applegate, 1965; Welton, 1979; Cappetta, 1987; Com- C A A RR R C Discoaster brouweri pagno, 1988).

C C R All the fossil material Casati & M. Tuscan El- Discoaster surculus (S. Zanaga

asmobranchCollection, Scandicci: CZ-TES 2 and S. Casati R R R RR R Discoaster pentaradiatus & M. Zanaga Tuscan Otolith Collection, Scandicci: CZ- R R Discoaster tamalis TOS 1) is deposited in the collectionsofthe “Centro civico

R R Ofelia Scandicci exhibitionthanks to Helicosphaerakamptneri/carterikamptneri/carteri Mangini, (permanent the “Gruppo AVIS Mineralogia Paleontologia Scandicci”). Cc R Helicosphaera sellii Address: Piazza Vittorio Veneto n.l, 50010 Badia a Set-

R Lanternites minutus timo, Scandicci, Firenze, Italy.

F Pontosphaera multipora

R Pontosphaera spp. Class Chondrichthyes Cc c cC cC cC Pseudoemiliania lacunosa Subclass Elasmobranchii

R Reticulofenestra pseudoumbilica (>7 pm) Cohort Euselachii Superorder Squalomorphii Compagno, 1973 C Vv | V V A A Reticulofenestra sp.sp, (<4 pm) Order HexanchiformesBuen, 1926 rTR Sphenolithusabies Family Chlamydoselachidae Garman, 1884

R Genus Garman, 1884 UmbiticosphaeraUmbilicosphaera spp. Chlamydoselachus

Table chart. 1. Nannofossil range

abundant A = (1-10 specimens per field of view), Chlamydoselachus lawleyi Davis, 1887

C 2-10 fields of = common (1 specimen per view), Figures 5/1-7, 6/1-6, 7/1,2

F =few 11-100 fields of (1 specimen per view),

R = 101-1000 rare (1 specimen per fields of view),

V = abundant field of 1876 very (11-100 specimens per view). Incertae sedis Lawley, p. 87, pi. 1, figs 1, la-c.

1983 C. Davis - 1-3. lawleyi Pfeil, p. 72, pi. 35, figs

discoasters do taxonomic 1986 C. cf. - 134. Generally, not present problems lawleyi Davis Cigala Fulgosi, p.

although their usefulness in the Mediterranean has been

questioned because oftheir alleged low abundance (Rio et

Materialexamined — A hundred of which al., 1990). However, the occurrence of a Discoaster as- teeth, some are and/or lack semblage with complete specimens in all the analyzed complete only the cusplets, all other teeth are

1 samples is an index of good preservation and a reliable fragmentary (CZ-TES 2, -98).

assemblage.

Remarks — The dental of The presence of three important markers as D. pentaradia- morphology Chlamydoselachus

surculus and tamalis allow to which is herein studied for first tus, D. D. correlatethe upper lawleyi. the time on the part ofthe studied section with the interval defined at the basis ofthis copious material, matches that ofthe living C.

the last occurrence of surculus anguineus & Smith, 1933; Goto, top by (LO) D. and D. pen- (Gudger 1972, 1977, taradiatus(2.5 Ma, Figure 2). The absence of sphenoliths 1987,2004;Goto & Hashimoto, 1976; Pfeil 1983; Welton,

We able and the presence ofD. tamalis allow the correlationof the 1979). were not to study the teeth ofC. sp. A, an

section with the D. tamalis-Zoneabove the last occurrence undescribed living species from Namibia (Ebert, 1990, of Sphenolithus spp. (3.7 Ma, Figure 2). The rare occur- 2003; Compagno et al., 2005). This latter species, if valid,

not be rence ofD. tamalis in two ofthe lower samples (i.e. 4 and might easily distinguishable from C. anguineus, , at

5; Table 1) could indicate the extinction event of this spe- least in considerationof its exterior features, and it would

cies (2.8 Ma). On this basis the sampled section is attrib- differ only on the basis of some anatomic features such as

uted to the middle-latepart ofthe Piacenzian/earliest Ge- the internal structure of the chondrocranium, the vertebra,

lasian stage (Late Pliocene, Figure 2). and the pectoral-fin skeletal morphology, etc. -6-

Figure 3. 1 -4, Dried head of ChlamydoselachusanguineusGarman, 1884. Specimen fished off Sesimbra (Lisboa, Portugal) (F. Cigala

1: lateral 2: frontal Fulgosi private collection 1, 951). view, x 1.1; view, x 0.6; 3; dorsal view, x 0.56.; 4: ventral view, x 0.62. -7-

Figure 4. Jaw of Chlamydoselachus anguineusGarman, 1884. Female, TL 1768 mm, off Kahbara, Suruga Bay (Japan); fishing gear

150-180 2 1985 K. Y bottom gill nets, depth m, May (Dr. ano) (F. Cigala Fulgosi private collection 1,120),x 1; a: upper jaw (latero-

b: lower d: lower posterior); upper jaw (antero-lateral); c; jaw (commissural-latero-posterior); jaw (lateral); e: upper jaw (symphysial,

anterior); f: lower jaw (symphysial, anterior). -8-

5. 1-7. 1 antero-lateral tooth h: Figure Chlamydoselachus lawleyi Davis, 1887; ; (CZ-TES 2, 1); a; labial;b, d, f, oblique; c, g: lateral;e;

lingual view, x 4.73; 2; antero-lateral tooth (CZ-TES 2. 2); a: labial; b, d: lateral; c: lingual; e; basal view, x 4.6; 3: antero-lateral

tooth (CZ-TES 2, 3); a: labial;b, d: lateral; c: lingual; e: basal view, x 3.95; 4; latero-posterior tooth (CZ-TES 2,4); a: labial; b, d:

lateral; c: lingual; e: basal view, x 5.7; 5: antero-lateral tooth (CZ-TES 2, 5); a: labial; b, d: lateral; c: lingual; e; basal; f: occlusal

view, x 4.2; 6: antero-lateral tooth (CZ-TES 2,6); a: labial;b, d: lateral;c; lingual; e: occlusal view, x 4; 7: antero-lateral tooth (CZ-

d: TES 2); a: labial;b, lateral; c: lingual; e: basal view, x 4,6. -9-

As Davis first 1875 Notidanus - (1887) pointed out on the basis ofthe only primigenius Agassiz Lawley, p. 20, pi.

tooth illustrated by Lawley, and confirmedby Pfeil (1983) 1, figs 1-5. - 1875 Notidanus gigas Sismonda Lawley, 23, pi. 1,figs on the basis of fragments and/or isolated cusps, the only p.

difference 6, 6a. we can perceive and/or definerelates to the size

1875 Notidanus - recurvus Agassiz Lawley, p. 24, pi. 2, of the teeth that, in the fossil species are nearly always figs 1, la, b. larger. The majority of teeth found are actually twice (or 1875 Notidanus targionii Lawley, p. 27, pi. 2, figs 3,3a, b. one and a half times) the size of those of an adult female 1875 Notidanus meneghiniiLawley, p. 28, pi. 2, figs3,3a, from specimen Japan measuring 1768 mm in length (coll. b, 4, 4a-c.

F. 1 n. 120, The fossil Cigala Fulgosi, Figure 4). largest 1875 Notidanus d’anconae Lawley, p. 29, pi. 3, figs 1, la, tooth is antero-lateral tooth with an (Figure 7/1) extremely 2, 2a.

divergent mesial and distal the 1875 Notidanus cusps (tooth height along problematicus Lawley, p. 31,pi. 3, figs 3,

median axis the base of the = 4a. cusp including root 9 mm; 3a, 4,

1875 Notidanus length of the median cusp from the apex to the enameloid anomaleLawley, p. 32, pi. 3, figs 5,5a, b,

6a. = 6, base - 8 mm; maximum mesio-distal width of the root

1909 Notidanus - de 5.8 maximumwidth griseus (Bonnaterre) Stefano, p.585, mm; ofthe tooth between the extremi- pi. 18, figs 3-10. ties of the mesial distal and cusps =13.4 mm; mediancusp

1912 - Notidanus griseus (Bonnaterre) de Stefano, p. 54, angle = 40°). pi. 2, fig. 30. The dental morphology and morphometries of living C.

1977 - Hexanchus griseus (Bonnaterre) Landini,p. 96, pi, anguineus were thoroughly examined by Welton (1979) 1, figs 1-6, 8-9. and Pfeil and shall therefore limit ourselves to (1983) we 1996 Hexanchus - Mane griseus (Bonnaterre) et al., p. 26, just a few observations. When referring to isolated fossil pi. 2, figs 6-7. teeth in Welton found it to 2001 - particular, impossible assign Hexanchus sp. Purdy et al., p, 84, fig. 5. these either to the upperor the lowerjaw and to definewith

the thedental series. The fossil Material — any certainty position along examined Three teeth (CZ-TES 2, n. 121-

teeth have less regularly secondary cusplets more or as high 123).

as the living ones; in one specimen, which as a result is

considered there three The — anomalous, are (Figure 5/2). Remarks Undoubtedly, the three teeth that were found

teeth of C. have and lawleyi regularly relatively sharp sig- belong to Hexanchus griseus, a species whose dentition, as

moid the of illustra- various cusps (confirming accuracy Lawley’s palaeontologists have observed, presents not only The characteristic be observed in the dried tion). same can ontogenetic and gynandric heterodonty (Purdy et al., head of from whose is inde- a living specimen Portugal sex 2001), but also a wide individual variability. One of the terminate coll. (Figure 3, F. 1 n. while Cigala Fulgosi, 951), teeth (Figure 7/6), which appears rather short even though in the jaw of the female specimen from mentioned Japan lacking the distal extremity, would appear to be the upper

above 4), the are robust and left of (Figure cusps slightly more eight tooth the symphysis. The other two, one of

less It is that this resemblance between the which is shown here sigmoid. likely (Figure 7/7), are typical lower jaw

fossils (in which this characteristic is constant), and the antero-lateral teeth; the illustrated tooth, relatively short be ascribed Portuguese specimen, cannot to a gynandric and high, is probably the first of the symphysis. Landini but the from close heterodonty to origin geographically (1977, pi. 12, figs 1-6, 8-9) illustrated several teeth from if (even chronologically distant) regions. the Lawley collection.

The rare frilled sharks have dis- a circumglobal deep water Hexanchus griseus is a large-sized species linked to the tribution even though they are captured only sporadically bottom, and generally inhabiting both deep waters (Torton- and in localised These demersal very areas. essentially ese, 1956)(upper slopes) and fairly deep waters (continen-

sharks seem to prefer the cold waters of and deep upwell- tal shelf to insular shelf). It appears that, at night, when ing regions. This is confirmed the catches in its by frequent chasing prey, this shark can reach the higher mass of the Suruga Bay, Japan in shallow waters (50-500), adjacent water, swimming even up to the surface. Furthermore, it

to deep regions (Kubota et al, 1991). Theirnocturnal as- be to find may possible young specimens “close inshore” in in the cent water column, almost to the surface, seems re- shallow waters 1984, This (Compagno, p. 20). species lated to The dietof the frilled sharks in- feeding. seems to seems to have migratory habits (Desbrosses, 1938) and in

clude other sharks and small teleosts which winter closer cephalopods, seems to come to the Sicilian coastline, reach-

to swallow in the snakes. they seem same manner as ing a depth of 300 metres (Arcidiacono, 1931).

Order Echinorhiniformes Buen, 1926

Hexanchidae 1851 Family Gray, Family EchinorhinidaeGill, 1862

Genus Hexanchus Rafinesquc, 1810 Genus Echinorhinus Blainville, 1816

Hexanchus griseus (Bonnaterre, 1788) Echinorhinus richiardiiLawley, 1876 7 Figures 7/6, Figures 7/3-5 - 10-

Figure 6. 1-6. Chlamydoselachus lawleyi Davis, 1887; 1: antero-lateral tooth (CZ-TES 2, 8); a, labial;b,d, lateral; c, lingual; e, basal

view, x 4.5; 2: antero-lateral tooth a; labial; b, lateral; (CZ-TES 2,9); d: c: lingual; e: basal view, x 5; 3: antero-lateral tooth (CZ-TES

2, labial; d: lateral; 7.3; 4: antero-lateral 10); a: b, c: lingual view, x tooth (CZ-TES 2, 11); a: labial;b, d: lateral; c: lingual; e: basal antero-lateral view, x 5.5; 5: tooth (CZ-TES 2,12); a: labial;b, d: lateral;c: lingual; e: basal view, x5.1; 6: posterior tooth (CZ-TES

2, 13); a: labial; b, d: lateral; c: lingualview, x 10.9. - 11 -

Figure 7.

1-2. Davis, 1887; 1: anterolateral tooth Chlamydoselachus lawleyi very large (CZ-TES 2, 14); a; labial;b: lingual-oblique view, x 3,45;

2: aberrant antero-lateral tooth (CZ-TES 2, 15); a: labial; b: lateral; c: basal view, x 2,6.

3-5. Echinorhinus richiardii Lawley, 1876; 3: antero-lateral tooth (CZ-TES 2, 124); a: labial; b: lingual view, x 2.2; 4: antero-lateral

tooth 2, a: b: labial view, x 2.8; (CZ-TES 125); lingual; 5: antero-lateral tooth (CZ-TES 2, 126); a: lingual; b: labial view, x 1.8.

6-7. Hexanchus 1810;6; left lateral tooth griseusRafmesque, upper (probably, the eighth from the symphysis) (CZ-TES 2,121); a: lin-

gual; b: labial view, x 1.13; 7: rightlower antero-lateral tooth (probably, the first) (CZ-TES 2,122); a: labial;b; lingualview, xO. 87. - 12-

8. cf. Centrophorus granulosus (Bloch & Schneider, 1801); right lower lateral tooth (CZ-TES 2, 130); a: lingual; b: labial view, x 4.2.

9-11. rostral rostral 9: 10: 11: Pristiophorus sp.. spines or teeth; (CZ-TES 2, 312), x 1.7; (CZ-TES 2, 313), x 1.6; (CZ-TES 2, 314), x

1.77,

12-16. antero-lateral 12: Squatina sp., teeth; (CZ-TES 2,326); a: lingual; b: labial view, x 1.88; 13: (CZ-TES 2,327); a: lingual; b: labial

view, x 2.47; 14: (CZ-TES 2,328), view, x 2.74; 15: (CZ-TES lingual 2, 329); lingual view, x 2,8; 16: (CZ-TES 2,330); labial view,

x 2.7.

17-19. 17: anterior tooth the labial Isurus oxyrinchus Rafinesque, 1810; right upper (probably first) (CZ-TES 2,342); a; lingual; b: view,

x 1.16; 18: left lower lateral tooth (CZ-TES 2, 343); a: lingual; b: labial view, x 0,87; 19; right lower anterior tooth (probably the

first) (CZ-TES 2, 344); a; lingual; b: labial view, x 1.2.

20-21 20; lower anterior tooth b: labial 1.2 21: Alopias superciliosus (Lowe, 1840); right (CZ-TES 2,339); a: lingual; view, x right up-

anterior lateral labial per or tooth (CZ-TES 2, 340); a: lingual; b: view, x 1.7.

anterior tooth the b: labial 22. Carcharias taurus Rafinesque, 1810; left upper (probably third) (CZ-TES 2, 335); a: lingual; view, x 1.

23. left lateral tooth b: labial 1.3. Odontaspisferox (Risso, 1810); upper (CZ-TES 2, 337); a: lingual; view, x

24-25. Prionace glauca (Linnaeus, 1758); 24: right lower lateral tooth (4th or 5th) (CZ-TES 2, 353); a: lingual; b: labial view, x 1.83;

25: left lateral tooth b: labial 2. upper (CZ-TES 2, 354); a: lingual; view, x

26, 28. Carcharhinus cf. 26: left tooth plumbeus (Nardo, 1827); upper (second or third) (CZ-TES 2,349); a; lingual; b: labial view, x 1.

28: left tooth 8th labial 2; upper (7th or ) (CZ-TES 2, 350); a; lingual; b; view, x 1.79.

27. Carcharhinus cf. perezi (Poey, 1876); left lower lateral tooth (6th or 7th) (CZ-TES 2, 352); a: labial; b: lingual view, x 1.68.

29. Dalatias licha (Bonnaterre, 1788); lower antero-lateral tooth (CZ-TES 2, 311); a: lingual; b: labial view, x 1.9.

1876 Echinorhinus richiardii Lawley, p, 41, pi. 1, figs 8, West Pacific and New Zealand), this species is most

8 a-b, 2, 6, 6 a-d. pi. figs probably a direct descendent of E. richiardii.

1977 Echinorhinus brucus (Bonnaterre) - Landini, p. Echinorhinusbrucus is a sluggish bottom shark, primar- 120, pi. 5, fig. 2 a-b. ily living in deep water (Compagno, 1984). Tortonese

(1956) claims that it is widespread in the Atlantic be-

Materialexamined — Five more or less complete teeth, tween 400 and 900 m and usually rare in the Mediterra- the largest of which has a low and sub-rectangular root nean. It is locally abundantand captured also in shallow 1.6 cm long. All the other teeth are approximately 1 cm water along the Senegal coast (6 specimens fished by line long, CZ-TES 2, 124-129. in a single pirogue, Cadenat & Blache, 1981).

Remarks — Echinorhinus teeth, usually rare, are fairly

well-represented in the studiedmaterial. As described by Order Squaliformes Goodrich, 1909 various authors ( e.g. Garrick, 1960; Bass et al., 1976; Family Centrophoridae Bleeker, 1859 Welton, 1979; Cadenat & Blache, 1981; Pfeil, 1983; Genus Centrophorus Muller & Henle, 1837 Purdy et al, 2001) the teeth ofthis genus have a hexan-

choid morphology and an anaulacorhizic vascularization.

Apart from the marked ontogenetic heterodonty (the teeth Centrophorus granulosus (Bloch & Schneider, 1801) of the young are monocuspidate), the various types of Figure 7/8 heterodonty are very feeble. As usual in sharks, it is dif- ficult to clearly separate fossil forms from living ones. - 1876 Acanthias major Agassiz Lawley, p. 40, pi. 1,

et al. noticed the of the Purdy (2001) variability typical figs 19, 19b.

few dentitionsofE. brucus that have been in - illustrated, 1909 Acanthias vulgaris Risso de Stefano, p. 594, p 1. particular in the more or less convex mesial cutting edge 7. figs 23, 24.

1972 & - of the main cusp and in its obliquity. Furthermore, it is Centrophorus granulosus (Bloch Schneider)

Ledoux, 145, 5, 6. not easy to distinguish the isolated teeth of E. brucus p. figs

1977 granulosus & - from those of E. cookei Pietschmann, 1928, due to the Centrophorus (Bloch Schneider) 4. Landini, p. 119, fig. scarcity of illustrated material (see also Pfeil, 1983).

The two living species, which are apparently similarand

Material examined — 180 the from the likely to have been confused in the past, actually differin teeth, majority the size and morphology ofskin denticles. The few den- lower jaw, CZ-TES 2, 130-310. ticles illustrated by Lawley (1876, pi. 2, figs 6c, d ) and

Remarks — There is subtle distinc- assigned to E. richiardiiare similar, or even the same, as actually only a very

tion between the teeth of Mediterranean those belonging to E. brucus. Pliocene specimens seem Centrophorus (Bloch & Schneider, 1801) and other Cen- to differvery littlefrom the latterspecies, the only differ- granulosus

C. ence being the larger size of the mesial cusplet close to trophorus species (i.e. i squamosus (Bonnaterre, 1788),

main the is still and E. C. lusitanicus & Capello, 1864,and C. niaukang the cusp. Clearly problem open Bocage in the Atlantic less close richiardii can be considered a species inquirenda (Wel- Teng, 1959) living more or to

identification at least has be based on ton, 1979),in the absence ofmore adequatecomparative Gibraltar; an to several teeth. Ledoux dealt with various material. Ifwe consider the geographicdistributionofE. (1970) already brucus (mainly East Atlantic and Mediterranean,but also types of heterodonty in C.granulosus (Bloch & Schnei- -13 -

der, 1801) and C. (Bonnaterre, con- waters more metres and squamosus 1788), than 200 sea depth it was cap- lusitanicus sidering C. Bocage & Capello, 1864 a form tured up to 1800 metres depth. Tortonese (1956) ob-

of C. and the or subspecies granulosus omitting large served that it is very rare in the Adriatic but present as a oceanic species C. niaukang Teng, 1959.On the basis of fossil in the Plioceneof the Adriatic Basin.

teeth and skin denticles Cigala Fulgosi (not illustrated,

1986, 1996) recognised the oceanic deep water species

C. in the squamosus early Zanclean (lower part of the Order Pristiophoriformes Berg, 1958

MPL2 biozone) of the Mediterranean palaeo-area. An Family Pristiophoridae Bleeker, 1859

analysis of approximately fifty jaws of C. granulosus Genus Pristiophorus Muller& Henle, 1837

(coll. F.C.F., from Sicily channel) suggests some varia-

tion in the shape of the root edge and in cusp serration,

which here to be more or less marked, but appears usu- Pristiophorus sp.

ally finerin C. squamosus (Ledoux, 1970). Furthermore, Figures 7/9-11

the teeth are rather small and of the same size as those

? found in most adult specimens from the Mediterranean 1876 Sphyraena Lawley, p. 87, pi. 1, figs 2, 2a,b.

(approximately 0.90-1 m TL, Cigala Fulgosi, pers. ob-

serv. ofhundredsof trawledoff In the specimens Sicily). Materialexamined — 14rostral spines (CZ-TES2,312- Atlantic (Madeira, Senegal) sexual maturity is reached at 325). size & a larger (1300 mm TL) (Cadenat Blache, 1981)

and we that the maximum size reaches at least

presume Remarks — This genus, known since the Late Creta- and half one metre a (Compagno, 1984). ceous, Late Santonian of Lebanon [i.e. Pristiophorus Like other species ofthis genus, C. granulosus often fre- tumidens (Woodward, 1932)is usually represented in the continental 400-800 quents slopes (upper slopes, m sea fossil record by isolated rostral spines (or saw teeth or

depth) and has, as an upper limit, the marginal shelf rostral “rostral teeth; spine” was used by Latham (1793)

(100- 200 m sea depth). both for Pristis and Pristiophorus). Rostral spines (e.g.

figs 7/9, 10) of approximately the same length have, in

the basal part ofthe crown, several short and sharp plicae Family DalatiidaeGray, 1851 which in other rostral spines of Pliocene Pristiophorus

Genus Dalatias Rafinesque, 1810 dense crenulation appear as a (Cigala Fulgosi, pers. ob-

serv.). Lawley (1876, pi. 1, figs 2, 2a, 2b) illustrated and

described two rostral spines foundin the Pliocene of Or-

Dalatias licha (Bonnaterre, 1788) ciano (Pisa, Tuscany, Italy), withoutrealising their na- Figure 7/29 ture. His figures, if observed through a magnifying lens,

present the same type of plicae. An undescribed deep 1876 Scymnus majori Lawley, p. 38, pl.l, figs 17, 17a, which water species, appears rather small in size and 17b. rostral represented by slight spines with some long pli- 1876 Centrina bassanii Lawley, p. 39, pi. 1, figs 18, cae, that might belong to P. schroederi Springer & 18a-e. Bullis, 1960) (Bahamas sawshark), was reported by Ci- 1909 - de Scymnus lichia (Cuvier) Stefano, p. 591, pi. gala Fulgosi (1986) as belonging to a psychrospheric 18, figs 16, 17. shark fauna (Cigala Fulgosi, 1996) from the Early Plio- 1977 licha - Scymnorhinus (Bonnaterre) Landini, p. of the MediterraneanBasin. 120, pi. 1, figs 20-24. cene

1986 Dalatias licha — Two short and flexed rostral (Bonnaterre) Cigala Fulgosi, p. spines (Figure 7/11) are

135, 137, fig. 2. similar to the kind of spines found in the region beneath

the eye ofPristiophorus species. The fossil rostral spines

Material — examined 1 tooth (CZ-TES 2, 311), were compared to those of three common living species

of Pristiophorus [viz. P. cirratus (Latham, 1794), P.

Remarks — Lowerjaw tooth froma specimen ofabout 1 nudipinnis Gunther,1870, and P. japonicus Gunther, in metre collected a thousand” 1870, which do not differ length. Having “certainly apparently among each other,

(“certamente un 38 and migliaio, Lawley (1876, p. “Orciano, which are different from the fossils, due to the base

Volterra, notedthat the teethofthis rare oftheir crown from smooth Siena”) species, being to very slightly crenu-

in the of Castelnuovo lated. Other undescribed in deposit Berardenga Scalo, are very living species are present in the Plioceneof less restricted common or frequent Tuscany. more or regions of Australia, the Indian

As summarised by Bigelow & Schroeder (1948), Com- Ocean and the Philippines (Last& Stevens; Compagno et

and et al. Dalatias al pagno (1984) Compagno (2005), ., 2005).

licha is With a widely ranging species locally abundant but not the exception ofP. nudipinnis which seems to in- ,

schooling. habit the continental shelves, the majority of living is It present in warm-temperateand tropical waters up to Pristiophorus species is found in the upper slope regions,

the North Scotland Irish Sea, and the Atlantic slope, around 500-600 metres sea depth, while P. schroederi,

South Australia and This which is the is from Africa, Japan. species prefers deepest species, reported 438-952m - 14-

sea depth, living on or near the bottom (Compagno etai, those of the living species, Carcharias taurus, of which

2005). six jaws were examined. One of these comes from the

Channel F.C.F. Sicily (coll. 1 n. 24, 1978)two from Ro-

cha, Punta del Diablo, Uruguay (coll. F.C.F. 1 n. 25 &

1973 Superorder Squatinomorphii Compagno, 112 ) and threefrom Madagascar (coll. F.C.F. 1 n. 653,

Order 1926 606 The Squatiniformes Buen, 927, ). dentitionof this species has been repeat-

Family Squatinidae Bonaparte, 1838 edly studied in order to determineboth the dentaltermi-

Dumeril, 1806 Squatina nology Ce.g. Applegate, 1965)andthe morpho-functional

aspects (Taniuchi, 1970; Luciforaetal., 2001). The teeth

often fine present plicae or striae which appear more de-

the basal halfof the Squatina sp. veloped on crown lingual face. The

Figures 7/12-16 illustrated tooth (Figure 7/22) seems to be a third upper

leftanterior tooth. The other, rather small, with an almost

1876 Squatina d’anconaiLawley, 37, pi. figs 16, and low p. 1, straight cusp and the base ofthe labial face pro- 16a-e. truding over the root, is a lower posterior tooth. 1909 Centrina salvianii Risso - de Stefano, 590, pi. p. This species, whichhas always been rare in the Mediter- 17, figs 27, 28. ranean (Doderlein, in: Tortonese, 1956, p. 103), could

1977 - Squatina squatina (Linnaeus) Landini, p. 121,pi. even have disappeared (apparently the last specimen, 5, fig. 6. coll. F.C.F. 1 n. 24, was collected by Cigala Fulgosi in

Mazara del Vallo, Sicily in 1978) and, if still present, is Material examined— Nine teeth (CZ-TES 2, 326-334). very rare along the North African coasts (Algeria and

Tunisia waters). It concerns a species with a wide distri- Remarks — It is difficult to identify the various species bution (Bass et al., 1975a; Cadenat & Blache, 1981; of Squatina on the basis of teeth alone. Even ifit were Compagno, 2001), in both coastal temperate-warm and possible, the now available material is certainly insuffi- tropical waters, and on continental shelves. It can be cient. Of fossil the nine teeth found, five are illustrated found in a variety of coastal habitats along sandy or (Figures 7/12-16). rocky beaches in shallow bays and reefs and submarine Squatina occurs today in the Mediterraneanwith three caves (Compagno, 2001, p. 60). It is a piscivorous spe- species: S. squatina (Linnaeus, 1758), S. aculeata Dume- cies (it eats various species of teleosts and small sharks) ril, 1829, and S. oculata Bonaparte, 1840, which fur- and be found alone in can or large aggregations or in thermore have a respective and different distributionin schools 2001, during breeding (Compagno, p. 60). the Eastern Atlantic (i.e. the region W of Gibraltar-the

Lusitanian-boreal region for the first mentioned species and from the Mauritanianregion to Angola, for the two Genus Odontaspis Agassiz, 1838 other species). Even the bathymetry appears to follow this separation, as S. squatina is the species that stays closest the while the to coasts, others live to a sea depth Odontaspis ferox (Risso, 1810) of approximately 500 metres. Indirectly, when the fossil Figure 7/23 assemblage indicates deep waters (e.g. upper slope) and the Squatina teeth are autochthonous (“not reworked”)

1912 - de Odontaspisferox(Risso) Stefano, p. 47, pi. 1, exclude first we can tentatively the species. fig. 17, pi 2, figs 11, 12.

1977 - Odontaspisferox (Risso) Landini, p. 103, pi. 1,

figs 13-17. 1973 1996 - Mane Superorder Galeomorphii Compagno, Odontaspis ferox (Risso) et al., p. 25, pi.

Order Lamniformes Berg, 1958 1, fig- 17.

Family Odontaspididae Muller & Henle, 1839 2001 Odontaspisferox (Risso) - Aguilera & de Aguil-

6/19. Genus Carcharias Rafinesque, 1810 era, p. 740, figs 6/18,

2006 - Suarez Odontaspisferox (Risso) etal., p. 12, figs

4j-k.

Carcharias taurus Rafinesque, 1810 — Material examined Two teeth (CZ-TES 2, 337, 338). Figure 7/22

— Remarks The only complete tooth (Figure 7/23) is 1977 Odontaspis (Synodontaspis) acutissima Agassiz - identical to those of the The tooth 10-12. living species. mor- Landini,p. 101,pf Lfigs phology of this species was with those of 1996 Odontaspis taurus (Rafinesque) - Mane et al., p. compared twelve Recent from 24, pi. l.figs 17-21. jaws, two the Mediterranean, viz. a male young (TL 215 cm) and a newborn female (TL 119

Material examined — Two teeth, CZ-TES 2, 335, 336. cm), and ten from the Indian Ocean, Madagascar (coll.

The individual F.C.F.). variationsfound agree with those

Remarks — The teeth be from described in two cannot distinguished literature. e.g. Cadenat & Blache (1981, fig- - 15 -

More less and 94). or high sharp teeth might represent a position ofthe fossil teeth in the jaw. Because ofthe pro-

slight sexual dimorphism. Someanomalies occur regard- nounced sexual dimorphism occurring in this species orientation ing the or the flexion ofthe (Bass et cusps. al., 1975a, p. 39, pi. 12; Gruber & Compagno, The here illustrated tooth (Figure 7/23), which more than 1981,figs 7,8; Cigala Fulgosi, 1983, pis 2-4; 1988,pi. 1, of the first likely represents one upper left latero- figs 1-3), the two fossil teeth can be attributed to male

teeth, has, as usual, three small lateral adult posterior cusplets specimens. In males the cusps are higher and

and the base of the labial face of the several at crown sharper (narrow at the base), with a convex labial face of

short and thick plicae. Its main cusp demonstrates dis- the crown. Because of the wide and U-shaped root, and tinct abrasion. Similar ofthe main strong wear cusp was the cusp, which is slightly bent towards the corner ofthe

also observed on the teeth of two or three of the more mouth, the first fossil tooth (Figure 7/20) might be the external series least (at two functional series) in three of second lower right tooth beginning from the symphysis.

the Recent The wear is common jaws. not only in almost The second (Figure 7/21), belonging to a smaller speci-

all but in of different teeth, occurs jaws a very size, men, with narrow and V-shaped root, and a relatively

the male from the Mediterranean bent namely young (F.C.F. cusp, might be the second upper right anterior or

coll.l n.630) and in one of the from one of the first larger specimens upper latero-posterior teeth. In the upper

Madagascar (F.C.F. coll. 1 n. 1023). This suggests that the jaw of this species, unlike A. vulpinus (Bonnaterre,

does not the of wear depend on longer persistence the 1788),the intermediatetoothis missing. The two anterior

teeth in functional in a position, as one might assume teeth, however, are disjunct from the latero-posteriors

We in of the because “ mature specimens. might suspect that, spite they are housed in the dental build’' (sensu functional and the of diet and Shimada, or “hollow” (sensu morphology type (fish 1998) Siverson, 1999, p. these sharks least cephalopods), feed, at occasionally, on 52, fig. 3), whichrepresents an important anatomic char-

the carcasses of other sharks, the of which is shagreen acter to define the heterodonty. A. superciliosus was re- abrasive. The other very toothlacks the root and the pro- cently signalled in the Mediterraneanby Compagno &

of the it could be of the first Gruber the basis of portions cusps suggest one (1981) on a specimen caught in the

lateral teeth. Ionian Sea. Shortly after, Cigala Fulgosi (1983) notedits

This which is almost in wide-ranging species, circumglobal occurrence the Sicily Channel. The same author (Ci- and confined to deep waters (upper slopes), can swim gala Fulgosi, 1988)also describedit from the Plioceneof

close to drop-offs in coral reefs. Recently, large speci- Tuscany (central Italy).

of this have been observed mens species by eco-tourist A. superciliosus is an epipelagic species (Cadenat & Bla-

divers in the Mediterranean and offMal- (off Lebanon) che, 1981), virtually circumtropical, which can be found

Island It is in pelo (East Pacific) (Compagno, 2001). gener- coastal waters and over the continental shelves and

but can be ally rare, locally abundant . in the also in far from (e.g rarely deep seas land (Compagno, 2001,p. 85).

Sicily Channel; specimens are captured along the coasts

of Madagascar). As Compagno (2001) observed, the

teeth of this species are less robust in comparison to Family Cetorhinidae Gill, 1862

those ofCarcharias taurus and seem to indicatean adap- Genus Cetorhinus Blainville, 1816

tation to and soft a more homogeneous prey (small fish, squid and shrimps).

Cetorhinus maximus (Gunner, 1765)

1876 - Family Alopiidae Bonaparte, 1838 (emended) non det. Lawley, p. 89, pi. 1, figs 11, 1 la-e. - Genus 1810 1876 Hannoveria van Beneden Alopias Rafinesque, aurata Lawley, p. 44,

pi. 1, fig. 17c.

1977 Cetorhinus maximus Gunner - Landini, p. Ill,

figs 18, 19. Alopias superciliosus (Lowe, 1840)

Figures 7/20-21

Material examined — One fragmentary gill-raker (CZ-

TES 2, 1911 - de 341). Oxyrhina Spallanzani Bonaparte Stefano, p.

396, pi. 10, figs 20, 21.

Remarks — 1988 - The shark is a Alopias superciliosus (Lowe) Cigala Fulgosi, p. basking represented by single of Gill rakers 95, pi. 1, figs 1,2. fragment a gill-raker. are more commonly

2001 - in Alopias aff. superciliosus (Lowe) Aguilera & de preserved the fossil record than the less numerous

Aguilera, 740, figs 6/22, 23. p. teeth, because in this shark the gill-rakers are much more

numerous than teeth.On their own, these elements do not

Material examined — Two teeth (CZ-TES 2, 339, 340). allow for a specific attribution (van den Bosch, 1984)

Remarks — The teeth be from two cannot distinguished and the identificationwith the living species is made in- those of the living species, Alopias superciliosus. A directly here. The living Cetorhinusmaximus (Gunner,

with some Recent comparison jaws (coll. F.C.F.), pre- 1765), has a long and complicated taxonomic history,

senting some variability also the regarding ontogenetic which was recently synthesised by Compagno (2001),

heterodonty, enables us to locate, at least tentatively, the who suggested the possible existence ofisolatedpopula- - 16-

tions with potentially restricted genetic interchange. Carcharhinus cf. plumbeus (Nardo, 1827)

This is is often huge species highly migratory and seen at Figures 7/26, 28 the surface of the epipelagic zone in deep water above

the slopes. Sometimes, in spring, when zooplankton 1996 Carcharhinus (Prionodon) egertoni (Agassiz) - blooms it also into shallow coastal develop, moves wa- Mane et al., p. 20, pi. 1, figs 1-5, 7.

2001 Carcharhinus plumbeus (Nardo) - Purdy et al., ters (Compagno, 2001, p. 93). p. 154, fig. 56.

Family Lamnidae Muller& Henle, 1838 Material examined — Three teeth (CZ-TES 2,349,350,

Genus Isurus Rafinesque, 1810 351).

Remarks — The few Carcharhinus teeth found are insuf-

Isurus 1810 oxyrinchus Rafinesque, ficient for any detailed taxonomic assessment. Nonethe-

7/17-19 of Figures less, some them (Figures 7/26, 28) are quite likely to

belong to C. plumbeus or to the lineage of that species,

1881 desorii - 77, 2, Oxyrhina Agassiz Lawley, p. pi. the teeth of which are occasionally also found in classic figs 1,1 a-b, 2, 2a-b, 3, 3a-b, 4-6, pi. 3, figs 1, la-b, Pliocene sedimentsof Parma and Piacenza (Cigala Ful- 2, 2a-b, 3, 3a-b. gosi, pers. observ.). For a betterassessment, the compari- 1899 Oxyrhina cf. gomphodonMuller & Henle -Vinassa son was extended to other fossil material from the Plio- de Regny, p. 82, pi 2, fig. 10. cene of other places in Tuscany where Carcharhinus

1909 - de Oxyrhyna Spallanzani Bonaparte Stefano, p. teeth are fairly common. The following species were rec- 570, pi. 16, figs 3, 8, 10,11. ognised (pers. obs. F.C.C.): C. obscurus (besueur, 1818),

1912 - de Oxyrhina spallanzani Bonaparte Stefano, p. C. cf. (Gunther, 1870), C. cf. 49, pi, 2, fig. 19. brachyurus brevipinna

C. 1977 Isurus - (Muller & Henle, 1839),

When the in the 1839)). present paper was proof, paper

of Marsili (2007) was published, in which C. plumbeus,

Materialexamined — Seven teeth(CZ-TES 2,342-348). C. brachyurus. C. falciformis and C. perezi were recog-

nized in the Pliocene of Tuscany.

Remarks — The of the teeth matches al. morphology in de- Recently Purdy et (2001, pp. 154-155, figs 55, 56), tail those observed several Recent dried with the of on jaws (coll. dealing taxonomy Carcharhinus, pointed out

F.C.F.) from various geographical locations. Ofthe three that C. obscurus can be distinguished fromi C. plumbeus illustrated teeth the smallest two (Figures 7/17, 19) are a in having the mesial cutting edge of the upper tooth first lower right anterior tooth and a first upper right crown apically convex as well. Using this character to- tooth, respectively. Their size is that of the teeth of Re- gether with the finer serration we can identify some of cent jaws of young specimens with a total length (TL) the teeth figured by Mane et al. (1996, figs 1-5, 7) as between 120and 130 cm. The third illustrated tooth(fig. belonging to the C. plumbeus lineage. Two Tuscan teeth

7/18) and a fourth, smaller tooth, are likely to be two (Figures 7/26,28) have a serration which, even though it lower left lateral is is finer teeth. The last three teeth, lacking part only fairly marked, relatively than that foundin of the root, are probably all second loweranterior teeth, C. obscurus. The latter, common in the PlioceneofTus- two on the left and one on the right. The smallest tooth cany and in the “Lawley collection” (see Landini, 1977,

is called has a crown the mesial side ofwhich measures 7.4 mmin pi. 4, figs 8-25) usually C. egertoni (Agassiz,

full-term it difficult length and it used to belong to a embryo or a 1843). Actually, is to understand to which juvenile. species of the group C. obscurus, C. leucas (Valen-

The “mako” is an extremely active and strong shark, dis- ciennes, 1839), C. plumbeus or C. galapagensis tributedworldwide in the tropical-temperate beltoffshore (Snodgrass & Heller, 1905)the specimens illustrated by but also in littoral waters. This species which is Agassiz (1833-1843, pi. 36, figs 6, 7) might refer. epipelagic and highly migratory, following the move- The first tooth(Figure 7/26) is narrow at the base and the

mesial and distal ments ofwarm water masses polewards, can reach waters crown edges have a style typical of the

least 500 2001, Ill second or third left tooth ofF C. of which at metres deep (Compagno, p. ). upper plumbeus.

When young, it consumes mainly small fish and cephalo- several Recent jaws were observed. The second (Figure pods, while large adults can attackother vertebrates such 7/28), a 7th-8thupperleft toothfrom the symphysis, pre-

notch the mesial and distal as sea turtles, other sharks and, rarely, marine mammals sents a along cutting edges

(dolphins). which render it similar to a tooth of C. falciformis. A

closercomparison between the teeth ofthese two species,

however, indicates that at times there are similar notches

Order CarcharhiniformesCompagno, 1973 also in C. plumbeus and that serration at the base ofthe

Family Carcharhinidae Jordan & Evermann, 1896 crown beneath the two notches is larger in C. falciformis

Genus Carcharhinus Blainville, 1816 than in C. plumbeus. - 17-

8. Present ofthe teleost in the Figure day bathymetric range taxa represented Piacenzian-earliest Gelasian ofCastelnuovo Berardenga

Scalo (Siena, Central Italy).

This does not in the fossil tooth which has ? 2001 - occur a more Carcharhinus perezi (Poey) Purdy et al., p. 154, regular serration along the cutting edges. The third tooth fig. 35b.

(not illustrated) is an upper lateral tooth.

Carcharhinus Material examined — One plumbeus today seems to be the most com- tooth (CZ-TES 2, 352).

mon species ofCarcharhinus caught in the Mediterranean

(Cigala Fulgosi, pers. observ., in the Sicily Channel). This Remarks — A comparison with some jaws (F.C.F. coll.) of

distributed this widely species (see Garrick, 1982) seems lo- species from Brazil and other teeth from the Pliocene

cally more or less abundant in temperate-tropical waters, of Tuscany (work in progress) has ledto this tentative iden-

with a of habitats. Indeed, it is found in tification. The serration is and the great variety regular develops up to

coastal shallow waters, but even in deep waters offshore or base ofthe crown on the mesial and distal shoulders where

close oceanic reefs. it is it becomes finer. The tooth to Furthermore, common at em- proportions enable us to con-

bayments, harbours, river estuaries etc. (Compagno, 1984, sider it as a 6th-7thlower left tooth from the symphysis. p. 494). Living Carcharhinus perezi is a bottom coastal Atlantic

species ranging from North Carolinato Venezuela-Brazil,

Carcharhinus cf. perezi (Poey, 1876) common in the Caribbean near drop-offs on outer reefs to

least Figure 7/27 at 30 metres in depth (Compagno et al., 2005). - 18-

Taxa _nn %

Coelorinchus coelorhincus 55 17.5

Trachyrincus scabrus 97 31 Macrouridae49.7%

Nezumia sp. 2 0.6

BathygadusBathyqadust sp. 2 0.6

Merluccius merluccius 15 4.8

Phycis blennoides 11 3.5

Micromesistius poutassouooutassou 4 121.2

Hoplostethus pisanusDisarms 69 22.1 TrachichthyidaeTrachichthvidae 22.1%

Rhynchoconger pantanellii 1 0.3

Pseudophichthys sp. 3 0.9

Hoplobrotula orcianensis 6 1.9

Lophiussp.sp. 3 0.9 Ceratoscopelus maderensis 1 0.3

Electrona rissoi 2 0.6

Lobianchia aff.aff.dofleinidofleini 1 0.3 Myctophidae 12% HygophumHygopbum hygomii 10 323.2

Diaphus splendidus 20 6.46,4

Diaphus holti 2 0.6

Notoscopelus cf. resplendens 2 0.6

Pagellus erythrinuservthrinus 6 I 1.9

Table 2. Recovered with of total number. otoliths ( sagittae ) per teleost taxa percentages

Genus Prionace 1849 Cantor. even though it tolerates warmer regions and can be found

in deeper, tropical seas (80-220 m, 12-25° C in tropical

India, Compagno, 1984, p. 522).

Prionace glauca (Linnaeus, 1758)

Figures 7/24, 25

Otolith fauna

1876 19 Prionodonsubglaucus Lawley, p. (pars).

1900 Prionodon Muller & Henle - glaucus Seguenza, p. Of otoliths, 312 teleostean sagittae (CZ-TOS 1, 1-312), 507, pi. 6, figs 37, 37a. representing 20 species (Table 2), were collected directly 1977 Prionace glauca (Linneo) - Landini, p. 117, pi. 3, from the surface. For taxonomic identifications, we used figs 9, 10, pi. 4, figs 1-3. various authors {e.g., Koken, 1891; Bassoli, 1905; Pierag-

noli, 1919; Dieni, 1968; Anfossi & Mosna, 1969, 1972, Materialexamined — 22 teeth (CZ-TES 2, 353-374). 1976, 1979; Gaemers, 1976, 1978; Gaemers & Schwarz-

hans, 1982; Schwarzhans, 1978a, 1978b, 1980; Nolf, 1980, Remarks — The teeth of this species are fairly common in

1985, 1994, 1995; Nolf & Cappetta, 1988; Nolf et al, the outcrop, although clearly less common than those of 1998; Flarkonen, 1986; Landini et al., 1990; Brzobohaty, Centrophorus and Chlamydoselachus. 1978, 1995; Brzobohaty & Nolf, 1995,1996,2000; Smale

etai, 1995;Girone & Varola, 2001; Girone, 2003; Lloris Their size ranges between small and modest, which indi- et al., 2003; Campana, 2004). cates that we are dealing with sub-adult specimens. This The identified species and/or genera, also the extinct ones, also appears to be true for the rare materialof Prionace are summarised in Figure 8 together with the (partly inter- from Early-Late Pliocene sediments of northern Italy (Ci- values of the preted) bathymetric corresponding living spe- gala Fulgosi, pers. observ.). cies (compare Bini, 1967-1969; Tortonese, 1970, 1975; The figured specimens represent a 4th-5thlowerright tooth Wheeler, 1969; Nafpaktitis et al., 1977; Noli & Cappetta, (Figure 7/24) and one of the last upper left lateral tooth 1988; Cohen et al., 1990; Nolf& Brzobohaty, 1994; Niel- (Figure 7/25), respectively. The teeth not illustrated here sen et al., 1999; Lloris et al., 2003; McEachran & Fech- consist of five lower laterals with small narrow cusps be- helm, 1998,2005 left laterals, etc.). longing to immature specimens, three upper and an upper right lateral tooth.

Because of its circumglobal distribution, which concerns the whole temperate-tropical range, Compagno (1984, p. Paleoecology

522) considered this species the widest ranging chondrich-

it cold thyan. However, appears to prefer waters (7-16°C), Chondrichthyan fauna - 19-

The most significant species from a bathymetric point of Conclusions

view be appear to Chlamydoselachus lawleyi and Centro-

phorus cf. granulosus of which approximately 100 (28%) The fossil fish assemblage from Castelnuovo Berardenga

and 180 teeth the Scalo (51%) were found, respectively. In case yielded macrourids (rat-fishes, grenadiers), mycto- of C. cf. granulosus. these meristic values are influenced phids (lantern-fishes) and shark genera,such as Centropho-

the number of individuals this by high represented by rus and Chlamydoselachus, indicating a Pliocene (Piacen-

in waters zian earliest bathypelagic species deep (100-1200 m, locally to Gelasian) bathyal environment, probably

more frequent between 400-800 m) near or at the bottom. located at short distance from the coast. The high number

The of high density Chlamydoselachus-teeth, concentrated of teeth belonging to Chlamydoselachus, a genus which

in such small is since the a area, exceptional, living species accomplishes great vertical migrations at night (Kubota et

is considered relatively rare everywhere. Kubota et al. al., 1991), is exceptional and might perhaps be related to

(1991), however, stressed that this rarity is apparent (de- some kind of upwelling, withoutexcluding other possible

the pending on fishing techniques) and that at least locally causes or mechanisms, such as currents and sea bottom

this shark be 139 can frequent (e.g. specimens were cap- depressions or traps. Its teeth, generally larger than those of

turedin four time in for examina- the toothwidth 13.5 years Suruga Bay, Japan, living species (max. mm. Figure 7/1),

tionof theirstomach contents). Even so, we cannot exclude suggest an animal approximately 2.5 metres in length (the

the possibility that some particular mechanism actually known maximum length of living C. anguineus is 1.96 m).

have caused these numbers. In Similar of might exceptional Suruga teeth equivalent size were recently reported from

Bay (Kubota et al., 1991, fig. 1) the sea bottom profile is the Early-Middle Miocene Tomioka Group, Japan (Taka- and steep bathyal depths are present within short distance kuwa et al., 2001). On the basis of size, we identify the

from the coast. Some Chlamydoselachus specimens were teeth as C. lawleyi. a fossil taxon whichto date was known

between51 and 60 from Zanclean captured during night at depths metres times, now demonstrated to persist into the

et al. thus the vertical Piacenzian-earliest Gelasian of the (Kubota , 1991), demonstrating great MediterraneanBasin.

ofthis to wandering bathybenthic mesopelagic species. The presence of Chlamydoselachus demonstratesthe per-

that indicate environment Species may an upper bathyal sistence ofa sufficiently deep connectionwith the Atlantic

continental Hexanchus without of (upper slope to outer shelf) are gri- Ocean, excluding some type upwelling. The

seus, Dalatias licha, Echinorhinus richiardii, Alopias su- absence of deep wateroceanic shark taxa like Deania, Cen-

and perciliosus Odontaspis ferox. The only shark species trophorus squamosus, Scymnodon spp., Scymnodalatias

the of which with the presence seems to contrast previous and Mollisquama (work in progress) present in the Early

is Carcharhinus cf. which Zanclean lower of picture perezi. prefers a strictly {i.e. part MPL 2 foraminiferalbiozone; coastal environment (inshore bottom dwelling species). see Cigula Fulgosi, 1986, 1996) of other Mediterranean

However, various of batoids types {e.g. many Dasyatis localities may be caused by insufficient localdepths during

and and species, Myliobatis Pteromylaeus) Notorynchus, deposition, by collecting bias, but also, and more probably,

in the coastal and in other Tuscan lesser of dwelling region present by a degree open oceanic setting of the Piacen- Pliocene outcrops, are lacking. Essentially, the chondrich- zian- earliest Gelasian Mediterranean Basin (Cigala Ful-

enable to thyans us hypothesise an upper bathyal environ- gosi, 1996).

ment probably close to the coast, and the high density of

Chlamydoselachus could perhaps berelated to some sort of

upwelling (high productivity). Acknowledgements

We the directorofthe of are grateful to quarry Castelnuovo

Teleosteanfauna (otoliths) Berardenga Scalo, Dr. MauroCartocci, forallowing us free

access to theoutcrop. Thanks are due to Mrs. E. Masinifor

The otolithassemblage (Table 2) is dominatedby Macro- the drawings and to Dr. Alex Orlandini (Reggio Emilia) for uridae (49.7%) and to a certain degree by Myctophidae (12 the photographic work (i.e. Figures 3-7). This %). co-occurrence was described as typical of deep

habitats {e.g. Nolf, 1985; Nolf & Brzobohaty, 1994). In

particular, benthopelagic species such as Trachyrincus References

scabrus and Coelorinchus coelorhincus are linked to the

bottom sea and represent 31% and 17.5%, respectively, of Agassiz, L. 1833-1843. Recherches sur les poissons fossiles, 3.

Neuchatel: viii + 82 the sagitta’s collected. This is alsorelated to the high num- 390, pis.

O. & de D.R. ber of individuals and specific diversity in which the Aguilera, Aguilera, 2001. An exceptional coastal fish in the Caribbean macrourids upwelling assemblage Neogene. Jour- naturally occur in deep habitats (Marshall, nal ofPaleontology, 75 (3): 732-742. 1979). Another bathypelagic (400-600 m depth) species, Anfossi, G. & Mosna, S. 1969. Otoliti del bacino terziario Lig- Hoplostethus pisanus (Trachichthyidae), is also common ure-Piemontese (Tortoniano Miocene e superiore). Atti (22.1%) in our material. The highest frequency of the rep- dell'IstitutoGeologico dell’Universitd di Pavia, 20; 23-49,5 resented taxa seems to be foundin sea depths between 300 pis. and 500 m (Figure 8, 90%), thus indicating an upper Anfossi, G. & Mosna, S. 1972. Otoliti del Pliocene inferiore di bathyal environment. Lugagnano(Piacenza). Atti dell’Istituto Geologicodella Uni- -20-

versitd di 23: 90-118, 7 inedites de la tribu Acanthiana Pavia, pis. especes de Squalidae Gray,

Anfossi, G. & Mosna, S. 1976. Otoliti del Pliocene inferiore della qui frequentent les cotes du Portugal. Proceedings of the

Liguria occidentale. Atti dell'lstituto Geologico dell’Uni- ZoologicalSociety ofLonden, (2): 260-263.

versitd di 26: 3 Pavia, 15-29, pis. Bonini, M. 2002. Extension and compression in the northern

Anfossi, G, & Mosna, S. 1979. La faunaittiologica di Monteu Apennines (Italy) hinterland: Evidence from the Late Mio-

Roero (Alba, Italia NW), otoliti. Alii dell'lstitutoGeologico cene-Pliocene Siena-Radicofani Basin and relations with

dell’Universitd di Pavia, 27: 111-132, 4 pis. basement structures. Tectonics, 21(3): 1-33.

M.T. H. 2002. Selaciens Anilines, & Cappetta, du Cretace C.L. 1832-1841. della italica (Al- Bonaparte, Iconografia fauna per

bien-Maastrichtien) d’Angola. Palaeontographica, A, Paldo- le quattro classi degli Animali Vertebrati, 3, Pesci. Roma,

zoologie-Stratigraphie, 264(5-6): 85-146, 12 pis. Tipografia Salviucci: 74 pis

Applegate, S.P. 1965.Tooth terminology and variation in sharks Bonnaterre, J.P. 1788. Tableau encyclopedique et methodique

with reference the sand Carcharias des trois de la Paris: vi special to shark. taurus regnes nature. Ichthyologic, 1. + 215

Contributions in Los 102 Rafinesque. Science, Angeles Country pp., pis.

Museum, 86; 1-18. Bosch, M. van den 1984.Oligocene to recent Cetorhinidae (Ver-

F. 1931. La del Arcidiacono, pesca pesce vacca (Hexanchus gri- tebrata, basking sharks); problematical finds ofteeth, dermal

seus) nella marina di Bollettinodi Pesca, Piscicol- scales and Riposto. gill-rakers. Mededelingen van de Werkgroep voor

tura e 608-612. Tertiaire Idrobiologia,7(4): enKwartaire Geologic, 21(4): 205-232, 3 pis.

Augugliaro, A. 1999-2000, Studi geologico-tecnici inerenti al Brzobohaty, R. 1978. ZurontogenetischeEntwicklung der fossi-

progetto di coltivazionee ripristino della zona di espansione len hoplostethiden (Otolithen, Miozan). Acta Musei Mora-

della cava di argilla posta in localitd Castelnuovo Ber- viae, 63: 49-56, 1 pi,

ardenga Scalo (SI). Unpublished thesis, Universita degli Brzobohaty, R. 1995. Macrouridae (Teleostei, Otolithen) im Oli-

Studi di Siena. und der zentralen und ihre gozan Miozan Paratethys palaoge-

Bass, A.J., d’Aubrey, J.D. & Kistnasamy, N. 1975a. Sharks ofthe ographische Bedeutung, Jahrbuch der Geologischen Bunde-

east coast ofsouthern Africa, 4. The families Odontaspididae, sanstalt, A, 138(4): 615-637, 5 pis.

Scapanorhynchidae, Isuridae, Cetorhinidae,Alopiidae, Orec- Brzobohaty, R. & Nolf, D. 1995. Diaphus otoliths from the

tolobidae and Rhiniodontidae. Investigational Report European Oligocene (Myctophidae, Teleostei). Bulletin de V

Oceanographic Research Institute,Durban, South Africa, 39: Institut royal des Sciences naturelles de Belgique, Sciences

1-102, 15 pis. de la Terre, 65: 257-268.

Bass, A.J., d’Aubrey, J.D. & Kistnasamy, N. 1975b. Sharks ofthe Brzobohaty, R, & Nolf, D. 1996. Otolithes de myctophides (pois-

east coast of southern Africa, 5. The families Hexanchidae, sons teleosteens) des terrains tertiaires d’Europe: revision des

and Chlamydoselachidae, Heterodontidae, Pristiophoridae genres Benthosema,Hygophum,Lampadena,Notoscopeluset

Squatinidae. Investigational Report Oceanographic Research Symbolophorus. Bulletin de I’Insitut royal des Sciences

Institute, Durban, South Africa, 43: 1-50, 9 pis. naturelles de Belgique, Sciences de la Terre, 66; 151-176,8

Bass, A.J., d’Aubrey, J.D. & Kistnasamy, N. 1976. Sharks ofthe pis.

east coast of southern Africa, 6. The families Oxynotidae, Buen, F. de 1926. Catalogo ictiologico delMediterraneo espanol

Dalatiidae and Echinorhinidae. de Marruecos lo sobre de las Squalidae, Investigational y recopilando publicado peces

Research South del de Report Oceanographic Institute, Durban, Af- costas mediterraneo y proximas Atlantico (Mar Es-

rica, 45: 1-103, 11 Commission Internationale pis. pana). Madrid, pour

Bassoli, G.G. 1905. Otoliti fossil! terziari deU’Emilia. Rivista de la Mer 221 FExploration scientifique Mediterranee: pp.

Italiana di Paleontologia, 12: 36-60, 2 pis. Cadenat, J. 1956. Note d’ichthyologie ouest-africaine, 14. Re-

L.S. 1958. der und le Berg, System rezenten fossilen Fischartigen marques biologiques sur requin-sable Carcharias (Odon-

und Fische. VEB der Wissenschaften: 310 1810. Bulletin de Berlin, Verlag pp. taspis) taurus Rafinesque 1’I.F.A.N. XVIII,

Bianucci, G., &Landini, W. 2005.1 paleositi a vertebrati fossili A (4): 1249-1256.

della provincia di Pisa. Atti della Societd Toscana di Scienze Cadenat, J. & Blache, J. 1981. Requins de Mediterranee et

naturali, Memorie,A, 110: 1-21. d’Atlantique (plus particulierement de la cote occidentale

Bibron,G. 1839. In: Muller, J. & Henle, F.G.J. (eds), 1838-1841, d’Afrique). Cahiers ORSTOM, Coll. Faune Tropicale, 21,

der Berlin. 330 Systematische Beschreibung Plagiostomen. pp.

Bigelow, H.B. & Schroeder, W.C. 1948. Sharks. In: Tee-Van, J., Campana, S.E. 2004. Photographic Atlas ofFish Otoliths of the

Breder, C.M., Hildebrand, S.F., Parr. A.E., Schroeder, W.C. Northwest Atlantic Ocean. Ottawa, Ontario, NRC Research

Fishes the North Sears Foundation Press; 284 (eds). of Western Atlantic, pp.

for Marine Research, Yale University, 1(1): 59-576. Cantor, T.E. 1849. Catalogue of Malayan fishes. Journal ofthe

Bini, G. 1967-1969. Atlante dei delle Asiatic 18; 983-1443 461 pesci coste italiane, Roma, Society ofBengal, i-xiii, (xiii + pp,

Mondo Soramerso Ed. Calcutta).

Blainville, H.M.D. de 1816. Prodrome d’une distribution syste- Capetta, H. 1987. Chondrichthyes II, Mesozoic and Cenozoic

du animal. matique regne Bulletin de la Societe Philomatique Elasmobranchii. In: Schultze, H.P., Kuhn, O. (eds). Hand-

Paris, 8; 105-124. book of Paleoichthyology, V. 3B: 1-193.

Bleeker, P. 1859. Enumeratio specierum piscium hucusque in Cigala Fulgosi, F. 1983. First record of Alopias superciliosus

Archipelago indico observatarum. Acta Societatis Regiae Sci- (Lowe, 1840) in the Mediterranean,with notes onsome fossil

6: 1-276. entiarum Indo-Neerlandicae, species ofthe genus Alopias. Annali del Museo civico di Sto-

Bloch M.E., & Schneider, J.G. 1801.M.E. Blochii Sistema Ich- ria Naturale di Genova, 84: 211-229, 4 pis.

Post auctoris F. A thyologiaeiconibus ex illustratum. obitum opus Cigala Fulgosi, 1986. deep water elasmobranch fauna froma

inchoatum ahsolvit, correxit, interpolavit J. G. Schneider. Lower Pliocene outcropping (northern Italy). In: Uyeno, T.,

Berlin, Saxo; ix + 584 pp., 110 pis. Arai, R., Taniuchi,T. & Matsuura K. (eds.). Indo-Pacificfish

J.V.B. du & Brito F. de 1864. Sur Bocage, Capello, quelques Biology, Proceedings Second International Conference on -21 -

Indo-Pacific Fishes. Tokyo: 133-139. 7 pis.

Cigala Fulgosi, F. 1988. Additions to the Eocene and Pliocene Gaemers, P.A.M. 1978. Late Quaternary and Recent otoliths from

fish fauna of Evidence of cf. denticulatus Italy. Alopias Cap- the seas around southern Norway. Mededelingen van de

1981 in the Bartonian-Priabonianofthe Monte Piano petta, Werkgroep voor Tertiaire en Kwartaire Geologic, 15(3);

Marl (northern Apennines) and of A. superciliosus (Lowe, 101-117, 2 pis.

1840) in the Pliocene ofTuscany (Chondrichthyes, Alopii- Gaemers, P.A.M.& Schwarzhans,W. 1982. Fisch-Otolithen aus

dae). Tertiary Research, 10(2): 93-99, 1 pi. der Typuslokalitat der obermiozanen Sylt-Stufe (Morsum-

Cigala Fulgosi, F. 1996. Rare oceanic deep water squaloid sharks Kliff, Insel Sylt, Nordwestdeutschland). Leidse Geologische

from the lower Pliocene of the northern Apennines (Parma Mededelingen, 52: 119-177, 10 pis.

province, Italy). Bollettino dellaSocietd Paleontologica Ital- Garman, S. 1884. An extraordinary shark (Chlamydoselachus

iana,34(3): 301-322, 6 pis. anguineus). Bulletin of the Essex Institute, 16: 47-55.

Cohen, - D.M., Inada,T., Iwamoto,T. & Scialabba,N. 1990. FAO Garman, S. 1885. Chlamydoselachus anguineus Garman a liv-

Species Catalogue, 10. Gadiform fishes of the world (order ing species of cladodont shark. Bulletin of the Museum of

Gadiformes). An annotated and illustratedcatalogue of cods, Comparative Zoology, 12(1): 1-35, 20 pis.

and other fishes known to date. J.A.F. 1960. hakes, grenadiers gadiform Garrick, Studies on New Zealand Elasmobranchii.

FAO Fisheries 442 Part X. The Synopsis, 125(10): pp. genus Echinorhinus, with an account ofa second

Colalongo,M.L. & Sartoni, S. 1967. Globorotalia hirsuta aemil- species, E. cookei, Pietschmann, 1928, from New Zealand

iana nuova sottospecie cronologica del Pliocene in Italia. waters. Transactions of the Royal Society ofNew Zealand, 88

Giornale di Geologia, ser. 2, 34: 265-285. (1): 105-117, pi. 7.

L.J.V. 1973. of elasmo- J.A.F. 1982. Sharks of the Carcharhinus. NOAA Compagno, Interrelationships living Garrick, genus

branchs. In-. R.S. & C. Technical Greenwood, P.H., Miles, Patterson, Report NMFS, 445: 194 pp.

(eds.). Interrelationships offishes. Zoological Journal ofthe Gill, 1862. Analyticalsynopsis of the order of squali and revision

Linnean London, 53 15-61, 2 the of the Society of (suppl.): pis. of nomenclature genera: squalorum generum no-

L.J.V. 1984. FAO 4. Sharks of Compagno, Species Catalogue, vorum descriptiones diagnosticae. Annals of the Lyceum of

the world. An annotated and illustrated catalogue of shark Natural History of New York, 7(32): 367-413.

species known to date, 1. Hexanchiformes to Lamniformes. Girone, A. 2003. The Pleistocene bathyal teleostean fauna of

FAO Fisheries viii Synopsis, 125(4): +249 pp. Archi (southern Italy): palaeoecological and palaeobio-

Compagno,L.J.V. 1988. Sharks the Order Rivista Italiana di of Carcharhiniformes. geographicimplications. Paleontologia e

Princeton, New Jersey, Princeton University Press: xxii + 486 Stratigrafia, 109(1): 99-110, 1 pi.

35 pp., pis. Girone, A., & Varola, A. 2001. Fish otoliths from the Middle

Compagno, L.J.V. 2001. Sharks ofthe world. An annotated and Pleistocene deposits of Montalbano Jonico (southern Italy).

illustrated catalogue of shark species known to date, 2. Bull- Bollettino della Societd Paleontologica Italiana, 40(3): 431-

head, mackerel and carpet sharks (Heterodontiformes,Lamni- 443, 4 pis.

formes and Orectolobiformes). FAO Species Catalogue for Goodrich, E. S. 1909. Vertebrata craniata, (First fascicle: cyclo-

Rome, FAO: 269 Fishery Purposes, 1(2), pp. stomes and fishes). In: Lankester, R. (ed.). A treatise on Zo-

Compagno,L.J.V., Dando, M. & Fowler, S. 2005. A field guide ology, 9. A. & xvi + 518 London, C. Black: pp.

the sharks the world. Collins: 368 to of pp. Goto, M. 1972. Fossil chondrichthyan of Japan. Geological

Davis, J.W. 1887.Note on a fossil species ofChlamydoselachus. Magazine, 78(11); 585-600.

the the So- M. 1977. Studies the teeth of Proceedings of Scientific Meetings of Zoological Goto, on Chlamydoselachus an-

London, 542-544. ciety of (1887): guineus, a living archaic fish, 2. On the development ofthe

Desbrosses, P. 1938. Croissance et migrations du requin-griset, teeth and the dermal teeth. Japanese Journal oforal Biology,

Hexanchus griseus Raf. 1810. Revue des Travauxde TOffice 19: 159-175.

des Peches Maritimes, 11:41. Goto, M. 1987. Chlamydoselachus anguineus - a living clado- Dieni, 1968. Gli otoliti Pliocene I. del inferiore di Orosei dont shark. Report of the Japanese Group of Elasmobranch

(Sardegna). Memorie della Accademia Patavina di Scienze, Studies, 23: 11-13.

Lettere ed Arti, Classe di Scienze Matematiche Naturali, e Goto, M. (and The Japanese Club for Fossil Shark Tooth Re-

80: 243-284, 3 pis. search), 2004. Tooth remains of chlamydoselachian sharks

Dumeril, A.M.C. 1806. Zoologie analytique ou methode from Japan and their phylogeny and paleoecology. Earth Sci-

naturelle de classification des animaux, rendue plusfacile a ence(Chikyu Kagaku), 58; 361-374.

I’aide de tableaux Paris: xxii 344 M. & I. 1976. the synoptiques. + pp. Goto, Hashimoto, Studies on teeth ofChlamy- Dumeril, A.M.C. 1829. Cuvier, In: G. 1829. Le Regne Animal, doselachus anguineus, a living archaic fish, 1. On the mor-

distribue son servir de base a d’apres organisation, pour phology, structure and composition of the teeth. Japanese

I’histoire naturelle des animaux et d'introduction a Journal of oral Biology, 18: 362-377, 2 pis.

I’anatomie 2: i-xviii ftn. 394. J.E. 1851. comparee, + 1-532, 2, p. Gray, List of the Specimens ofFish in the Collection of

Ebert, D.A. 1990. The Taxonomy, Biogeography, and Biology of the British Museum, 1. Chondropterygii. London, British

Cow and Sharks Museum (Natural 160 frilled (Chondrichthyes: Hexanchiformes). History), pp.

Unpublished phylosophy doctor dissertation, Rhodes Univer- Gruber, S.H. & Compagno,L.J.V. 1981. Taxonomic status and

Grahamstown, South Africa: 308 sity, pp. biology ofthe bigeye thresher, Alopias superciliosus. Fishery

Ebert, D.A. 2003. Sharks, Rays, and Chimaeras of California. Bulletin, 79(3): 617-640. California Natural History Guides. Berkeley, University of Gudger, E.W. & Smith, B.G. 1933. The natural History of the

California Press: xiii 284 + pp. frilled Shark, Chlamydoselachus anguineus. The Bashford

Gaemers, P.A.M. 1976. New gadiform otoliths from the Tertiary Dean Memorial Volume,Archaic Fishes, 5. New York, The

of the North Sea Basin and a revision ofsome fossil and re- American Museum of Natural History: 243-330, 5 pis.

cent species. LeidseGeologische Mededelingen,49: 507-537, Gunner, J.E. 1765. Brugden (Squalus maximus), beskrivenen ved -22-

J.E. Gunnerus. Det Trondheim. Selskab Fines 2. 57 12 Skrifter, 3:33-49, pi de la Pesca, Roma, pp., pis.

2. Lourens, L.J., Antonarakou, A., Hilgen, F.J.. Hoof, A.A.M. van,

Gunther, A 1870. Catalogue of thefishes in the British Museum. Vergnaud-Grazzini, C. & Zachariasse, W.J. 1996. Evaluation

Catalogue ofthe Physostomi containing the families Gymno- of the Plio-Pleistocene astronomical timescale. Paleocean-

tidae,Symbranchidae, Muraenidae,Pegasidae and ofthe Lo- ography,11(4): 391-413.

phobranchii,Plectognathi, Dipnoi, Ganoidei,Chondroptery- Lourens, L.J., Hilgen,F., Shackleton, N.J., Laskar, J. & Wilson,

gii. Cyclostomata, Leptocardii in the collection ofthe Britisch D. 2004. The Neogene Period. In: Gradstein, F., Ogg, J. &

Museum, London, (Natural History), 8: i-xxv + 1-549. Smith A.G. (eds.). A Geologic Time Scale 2004. Cambridge

T. 1986. Guide the Otoliths the Fishes Harkonen, to of bony of University Press, U.K: 589 pp.

the Northeast Atlantic. Danbiu 256 R.T. 1840. A from the Rev. R.T. M.A. Hellerup, Denmark, ApS: Lowe, paper Lowe, de-

97 certain ofMadeiran and contain- pp, pis. scribing newspecies fishes,

Jordan, D.S. & Evermann,B.W. 1896-1900. The fishes ofNorth ing additional information relating tothose already described.

and Middle America, a descriptive catalogue ofthe species of Proceedings of the ZoologicalSociety ofLondon, 1840: 36-

fish-like vertebrates found in the waters of North America 39.

north ofthe isthmus of Panama. Bulletin ofthe U.S. national Lucifora, L.O., Menni, R.C. & Escalante, A.H. 2001. Analysis of

Museum, 47( 1)( 1896): i-ix,1-1240. dental insertion angles in the sand tiger shark, Carcharias tau-

Koken, E. 1891. Neue Untersuchungen an tertiaren Fisch- rus (Chondrichthyes: Lamniformes). Cybium, 25(1): 23-31.

Otolithen, 2. Zeitschrift der Deutschen geologischen GeselT Mane, R., Magrans, J. & Ferrer, E. 1996. Ictiologia fossil del

schaft, 43: 77-170, 10 pis. Plioce del Baix Llobregat, 2. Selacis pleurotremats. Batalle-

Kubota, T., Shiobara, Y. & Kubodera, T. 1991. Food habits of ria, 6: 19-33, 2 pis.

the frilled shark Chlamydoselachus anguineus collected from Manganelli, G., Benocci, A. & Spadini, V. 2006. The scientific

Suruga Bay, Central Japan.Nippon Suisan Gakkaishi, 57(1): bibliography ofRoberto Lawley (1818-1881)and his contri-

15-20. bution to the study offossil sharks. Archives ofNatural His-

W. “ittiodontoliti Landini, 1977. Revisione degli pliocenici” tory, 33(2): 267-281.

della collezione Lawley. PalaeontographiaItalica, 70 (n. ser. Marshall,N.B. 1979. Developments in deep-sea Biology. Poole

5 Blandford Press Ltd: 566 40): 92-134, pis. UK, pp.

G. & C. 2005. II di S. 2007. Revision the teeth of the Carcharhinus Landini, W., Bianucci, Sorbini, giacimento Marsili, of genus

Orciano (Pisa). In'. Bonfiglio, L.(ed.), Paleontologia dei Ver- (Elasmobranchii; Carcharhinidae) from the Pliocene of Tus-

tebrati in Italia. Evoluzione biologica, significato ambientale cany, Italy. Rivista Italiana di Paleontologiae Stratigrafia,

epaleogeografico, Memorie del Museo civico di Storia natu- 113(1): 79-95, 2 pis.

rale di Verona, 2. Sez. Scienze della Terra, 6: 180-182. McEachran, J.D. & Fechhelm, J.D. 1998. Fishes of the Gulf of

Landini, W., Menesini, E. & Ragaini, R. 1990. Paleocomunita a Mexico, I. Myxiniformes to Gasterosteiformes. Austin, Uni-

molluschi ed otoliti nel Pliocene di Castelfiorentino of Texas Press: 1112 (Firenze, versity pp.

Italia), Atti Societd Toscana di Scienze Natural! Memorie, McEachran, J.D, & Fechhelm, J.D. 2005. Fishes of the Gulf of

(A)97: 175-202. Mexico 2. Scorpaeniformes to Tetraodontiformes. Austin,

J. 1794. An the various of sawfish. of Texas Press: 1004 Latham, essay on species University pp.

Transactions of the Linnean Society of London, 2(25): 273- Muller, J. & Henle, F.G.J. 1837. Gattungen der Haifische und

282, 1 pi. Rochen, nach Hirer Arbeit: Ueber die Naturgeschichte der

Lawley, R. 1875. Monografia del Genere Notidanus rinvenuti Kndrpelfische. Sitzungsberichte der k. preussischen Akade-

alio Stato del Toscano. mie der 2: 111-118. fossile Pliocene subappennino Wissenshaften zu Berlin,

Stab. 4 J. & F.G.J. Firenze, Pellas: 34 pp, pis. Muller, Henle, 1839-41.Systematische Beschreibung

R. 1876. Nuovi studi ai ed altri der Veit & Lawley, sopra pesci vertebrati Plagiostomen. Co., Berlin, i-xxii + 1-200,60 pis

delle colline delFArte 1-28 in 29-102 in fossili toscane. Firenze, Tipografia [pp. published 1838,reset pp. 27-28, 1839,

della Stampa: 1-122, 5 pis. i-xxii + 103-200 in 1841].

Lawley, R. 1881. Studi comparativi sui pesci fossili coi viventi Nafpaktitis, B.G., Backus, R.H.. Craddock, J.E., Haedrich, R.L.,

dei generi Carcharodon, Oxyrhina e Galeocerdo. Pisa, Tip. Robison, B.H. & Karnella, C. 1977. Family Myctophidae. In:

Nistri 151 25 7. New Sears Foundation Marine &c., pp., pis. FWNA, Haven, for Re-

Ledoux, J.-C. 1970. Les dents des squalides de la Mediterranee search: 13-265,

occidentale et de 1’ Atlantiquenord-ouest africain. Vie et Mi- Nardo, G.D. 1827. Prodromus observationum et disquisitionum

lieu,A. Biologic Marine. 21(2-A): 309-362. Adriaticae ichthyologiae. Giornaledi Fisica, Chimica, Stnria

Ledoux, J.-C. 1972. Les Squalidae (Euselachii) miocenes des Naturale , Medicina, Arti, Pavia, 10: 22-40.

environs d’Avignon(Vaucluse). Documents des Laboratoires Nielsen, J.G., Cohen, D.M., Markle, D.F. & Robins, C.R. 1999.

de Geologic de la Facultedes Sciences deLyon, 52: 133-175. FAO species catalogue, 18. Ophidiiform fishes of the world

Lesueur, C.A. 1818. Description of several new species of north (order Ophidiiformes). An annotated and illustrated catalogue

american fishes. Journal of the Academy of Natural Sciences of pearlfishes, cusk-eels, brotulas and other ophidiiform

of Philadelphia, l(pt. 2): 222-235; 359-368, pis 10, II, fishes known to date. FAO Fisheries Synopsis, 18. Rome,

222-235 14[pp. published in May, pp. 359-368 in Sept.]. FAO: 94-135.

C. 1758. Naturae tria D. 1980. Etude Linneaus, Systema per Regna Naturae, Nolf, monographique des otolithes des Ophidi-

secundum iformes actuels revision des fossiles Teleo- Regnum animale, Classes, Ordines, Genera, Spe- et especes (Pisces,

Characteribus cies, cum differentiis Synonymis, Locis, editio stei). Mededelingen van de Werkgroep voor Tertiaire en

decima. Holmiae, Salvius: 824 Kwartaire 71-195, 20 pp. Geologic,Geologic, 17(2): pis.

Lloris, D., Matallanas,J. & Oliver, P. 2003. Merluzas del mundo Nolf, D. 1985. Otolithi Piscium. In: Schultze, H.P. (ed.). Hand-

(Familia Merlucciidae). Catalogo comentado e ilustrado de book of Palaeoichthyology, 10. Stuttgart & Berlin, Gustav

las merluzas conocidas. FAO de los Fischer: 145 Catalogo Especies para pp. -23-

D. 1995. Studies fossil Nolf, on otoliths. The state ofthe art. In: permost Gearle Siltstone (Cenomanian, Late Cretaceous) of

Secor, D.H., Dean, J.M. &Campana, S.E. (eds.). Recent De- Western Australia. Transactions ofthe Royal Society of Ed-

velopments in Fish Otolith Research. Columbia, University inburgh, Earth Sciences, 90: 49-66.

Press: 735 & of South Carolina pp. Smale, M., Watson, G., Hecht, T. 1995. Otolith atlas of south-

Nolf, D. &Brzobohaty, R. 1994. Fish otoliths as paleobathymet- ern African marine fishes. Ichthyological Monographsof the

ric indicators. In: Agusti, J. (ed). Paleontologia i Evolucio. J.L.B. Smith Institute 1: 253 149 of Ichthyology, pp., pis.

the IX Proceedings of Congress, Regional Committee on Smith, B.G. 1937. The anatomy ofthe frilled shark, Chlamydose-

MediterraneanNeogeneStratigraphy, Barcelona, Nov. 1990: lachus anguineusGarman. The Basford Dean Memorial Vol- 24-25 255-264. (1992), ume. Archaic Fishes, 6. New York, The American Museum

Nolf, D. & H. 1988. Otolithes Cappetta, depoissons pliocenes du ofNatural History: 331-520,7 pis.

Sud-Est de la France. Bulletin de I Tnstitut royal des Sciences Snodgras, R.E. & Heller, E. 1905. Papers fromthe Hopkins Stan-

naturelles de Belgique, 58: 209-271, 18 pis. ford Galapagosexpedition, 1898-1899. XVII. Shore fishes of

Nolf, D. & Girone, A, 2006. Otolithes de poissons du Pliocene the Revillagigedo, Clipperton, Cocos and Galapagos Island.

inferieur (Zancleen) des environs d’Alba (Piemont) et de la Proceedings of the Washington Academyof Sciences, 6: 333-

cote ligure. Rivista Piemontese di Storia Naturale, 27: 77- 427.

114, 10 pis. Springer, S. & Bullis, H.R. 1960. A new species of sawshark,

R. & A. 1998. Otolithes Nolf, D., Mane, Lopez, de poissons du Pristiophorus schroederi, from the Bahamas. Bulletin ofMa-

inferieur de de Barcelone. Pliocene Papiol, pres Palaeoverte- rine Science of the Gulf and Caribbean, 10(2); 241-254. brata, 2T(l-2): 1-17. Stefano, G. de 1909. Osservazioni sulla ittiofauna pliocenica di Pfeil,F.H. 1983. ZahnmorphologischeUntersuchungenan rezen- Orciano e San Quirico in Toscana. Bollettino della Societd

ten und fossilen Haien der OrdnungenChlamydoselachifor- geologica Italiana,28: 539-648, 5 pis.

mes und Echinorhiniformes. Palaeoichthyologica, 1: 1-315. Stefano.G.de 1911, Sui pesci pliocenici dell'imolese.Bollettino

Pieragnoli, L. 1919. Otoliti plioceniche della Toscana. Rivista dellaSocietd geologica Italiana,29: 381-402, 1 pi.

italiana di Paleontologia, 25:21-44. Stefano, G. de 1912. Appunti sulla ittiofauna fossile deU’Emilia

Pietschmann, V. 1928. Neue Fish-arten aus dem Pacifischen conservata nel Museo Geologico dell’Universitadi Parma.

Ozean. Anzeiger der Akademie der Wissenschaften in Wien, Bollettinodella Societd geologicaItaliana, 31: 35-78, 2 pis.

65: 297-298. Suarez, M.E., Encinas, A. & Ward, D. 2006. An Early Miocene

Poey, F. 1876. Enumeratio piscium cubensium. Anales de la elasmobranch fauna from the Navidad Formation, Central

Sociedad Espanola de Historia Natural, Madrid, 5:131-218. Chile, South America. Cainozoic Research, 4(1-2); 3-18. P. 1920. Ittiofauna fossile dell’Italia Principi, centrale. Bollettino Takakuwa, Y., Goto, M., Hasegawa, Y., Yamazawa, T., Taka-

della Societd 39: 1 Y. & M. 2001. Tooth GeologicaItaliana, 85-110, pi. yama, Shimizu, remains of Chlamy-

Purdy, R. W., Schneider, V.P., Applegate, S.P., McLellan, J.H., doselachus (Chondrichthyes, Elasmobranchii) from the Tomi-

Meyer, R.L. & Slaughter, B.H. 2001. The Neogene sharks, oka Group (Lower to Middle Miocene) in Tomioka and An-

and fishes from Lee Creek North naka Gunma rays, bony Mine, Aurora, City, prefecture, central Japan. Bulletin of the Carolina. In: Clayton E.R. & Bohaska, D.J. (eds.). Geology Gunma Museum Natural History, 5; 19-30, 1 pi.

and Paleontology of the Lee Creek Mine,North Carolina, 3. Taniuchi, T. 1970. Variation in the teeth ofthe sand shark, Odon-

Smithsonian Contributions to Paleobiology, 90: 71-202. taspis taurus (Rafinesque) taken from the East China Sea.

Rafinesque, C.S. 1810. Caratteri di alcuni nuovi Generi e nuove Japanese Journal of Ichthyology, 17(1): 37-44. di Animali della Specie e Piante Sicilia con varie Osservazi- Tortonese, E. 1956. Leptocardia, Ciclostomata,Selachii. Fauna

oni i medesimi. Palermo, San 105 20 Calderini: viii sopra Filippo: pp., pis. dTtalia. Bologna, Ed. + 332 pp.

I. & G. 1990. Pliocene-Pleistocene Rio. D., Raffi, Villa, calcare- Tortonese, E. 1970. Osteichthyes (Pesciossei), 1. Fauna dTtalia.

nannofossil distribution in the western Mediter- Ed. 565 ous patterns Bologna, Calderini: pp.

ranean. In: Kastens K.A., Mascle J. etalfeds.). Proceedings Tortonese, E. 1975. Osteichthyes (Pesciossei), 2. Fauna dTtalia.

Ocean 107:513-533. Ed. 636 Drilling Programme, Scientific Result, Bologna, Calderini: pp.

Risso, A. 1810, Ichthyologic de Nice, ou histoire naturelle des Vinassa de Regny, P. 1899. Pesci neogenici del Bolognese. Ri-

poissons du department des Alpes Maritimes. Paris, F. vista Italiana di Paleontologia, 5: 79-84, 1 pi.

xxxvi 388 11 A. Asher & Schoell, + pp., pis. (reprinted Co. Ward, DJ. & Galea Bonavia, C. 2001. Additions to, andareview

the Amsterdam, 1966). of, Miocene shark and ray fauna of Malta. The Central

E. 1970. Otoliti del Robba, Tortoniano-tipo(Piemonte). Rivista MediterraneanNaturalist, 3(3): 131-146, 2 pis.

Italiana di Paleontologia,76(1): 89-172, 9 pis. Wheeler,A. 1969. Thefishes of the British Isles and North West

Schwarzhans, W. 1978a. Otolithen aus dem Unter-Pliozan Macmillan: 613 16 von Europe, London, pp., pis.

Siid-Sizilien und aus der Toscana. Berliner Geowissen- Welton, B.J. 1979. Late Cretaceous and Cenozoic Squalomorphii

schaftliche Abhandlungen,A, 8: 1-52, 13 pis. of the northwest Pacific Ocean. Berkeley, University ofCali-

Schwarzhans, W. 1978b. Otolith and ist for fornia 553 morphology usage (unpublished dissertation): pp.

higher systematical units, with special reference to the Myc- Woodward, A.S. 1932. A Cretaceous Pristiophorid shark. Annals

tophiformes s.l. Mededelingen van de Werkgroep voor Ter- and Magazineof Natural History, 10: 476-478, pi. 18.

tiaire en Kwartaire Geologic, 15(4): 167-185.

Seguenza, L. 1900.1 vertebral! fossil! della provincia di Messina,

1. Pesci. Bollettino della Societd Geologica Italiana, 19:

444-520, 2 pis.

Shimada,K. 1998. Dental homology in macrophagous lamniform

sharks. Journal of Vertebrate Paleontology, 18, (supplement

to n. 3, 7A) (abstract only).

Siverson, M. 1999. A lamniform shark from the new large up-