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110
Sicilian Cenozoic sharks from the collections of
the G. G. Gemmellaro Museum
Gli squali del Cenozoico di Sicilia
nelle collezioni del Museo “G. G. Gemmellaro”
ANTONIO CUSUMANO & CAROLINA DI PATTI
Museo Geologico “G. G. Gemmellaro” - Dipartimento di Geologia e Geodesia dell’Università di Palermo.
C.so Tukory n. 131 - 90134, Palermo.
E-mail: dipatti@unipa.it
AbstrAct
Among the several studies initiated by Gaetano Giorgio and Mariano Gemmellaro, some deal with osseus and cartilagineous shes. e nds fall
within a broader spectrum of collections and originate mainly from Sicilian palaeogenic, neogenic and quaternary soils.
is paper sets out to consider cartilagineous shes alone. It was G. G. Gemmellaro that prompted early classication of such nds in the mid-
19th century. At the beginning of the 20th century, Mariano Gemmellaro reviewed the material systematised by his father and moved onto index
that which was subsequently unearthed. Almost a century later, most of M. Gemmellaro’s cataloguing was found to be very consistent with only a
handful of updates in nomenclature needed.
e data retrieved was then being referred to as many as 40 species, seven of which had been determined by Gaetano Giorgio, one by Mariano.
However, on reviewing the pieces recently, the number of species was reduced to 18, three of which are still living.
e material dates to between the Middle Eocene and the Early Pleistocene, with greater concentrations from the Early and Middle Miocene.
One point worth of note is the common geolithological provenance of most nds: calcarenite and bio-calcarenite, some of which glauconitic,
suggesting low sea-levels; the presence of some open-water shark species in association to such geology makes it likely these were ideal hunting
areas.
Furthermore, deposits predominantly display large sized individuals together with higher than usual concentrations of Isurus, Carcharodon and
Odontaspis remains.
As to palaeoclimate and palaoecology, the majority of faunal assemblages considered – though by no means contemporary – derive from neritic
contexts, with water temperatures warmer than present and rich prey.
KEY WOrDs : Vertebrates (Chondrichthyes, Elasmobranchii), Sharks, Teeth, Cenozoic, Sicily, Systematic.
riAssuntO
Tra le innumerevoli ricerche arontate da Gaetano Giorgio prima, e da Mariano Gemmellaro poi, vi sono alcuni studi che riguardano pesci ossei e
cartilaginei. I reperti sono inseriti in collezioni più ampie e provengono principalmente dai terreni paleogenici, neogenici e quaternari della Sicilia.
In questo lavoro sono analizzati soltanto i reperti appartenenti ai pesci cartilaginei. Una prima classicazione di tali reperti era stata eettuata,
nella seconda metà del XIX secolo, da G. G. Gemmellaro. Agli inizi del XX secolo, Mariano Gemmellaro revisionava il materiale studiato dal padre
e classicava i reperti rinvenuti successivamente. A distanza di quasi un secolo, le determinazioni eettuate da M. Gemmellaro si sono rivelate in gran
parte corrette necessitando, soltanto in alcuni casi, di un aggiornamento della nomenclatura.
I reperti riferiti a ben 40 specie, di cui sette istituite da Gaetano Giorgio e una da Mariano Gemmellaro, dopo una recente revisione, si sono
ridotti a 18 specie, di cui 3 ancora viventi.
L’età del materiale analizzato nel presente lavoro va dall’Eocene medio al Pleistocene Inferiore, con maggiore abbondanza di ritrovamenti nel
Miocene inferiore e medio.
Un dato signicativo emerso è la provenienza, per la maggioranza dei reperti, da una litologia comune: calcareniti e/o biocalcareniti, talora
glauconitiche, che denotano l’esistenza di un mare poco profondo; la presenza di specie di squalo, tipiche di mare aperto, evidenzia come queste aree
dovessero rappresentare un territorio di caccia ideale.
Una ulteriore peculiarità è la prevalenza di esemplari di grossa taglia e il ritrovamento, con maggiore frequenza, di resti appartenenti ai generi
Isurus, Carcharodon e Odontaspis.
Quaderni del Museo Geologico Gemmellaro Volume 9 pp. 110-120 Anno 2006
111
Da un punto di vista paleoclimatico e paleoecologico, l’insieme delle faune considerate – pur non provenendo da livelli coevi – ha posto in
rilievo una netta prevalenza di forme sviluppatesi in habitat di tipo neritico, caratterizzati, nel complesso, da acque più calde di quelle attuali e ricche
di prede.
tErMini cHiAVE : Vertebrati (Condritti, Elasmobranchi), Squali, Denti, Cenozoico, Sicilia, Sistematica.
cOLLEctiOns’ HistOrY
Several cartilagineous sh teeth in excellent state of preservation are kept at Palermo University’s G. G. Gemmellaro
Geology Museum. ey essentially comprise of sharks, many fragmentary pieces as well as spine parts; 550 originate from
Sicilian rocks.
e nds hereby analysed belong to a number of collections. e authors chose to browse through these pieces
according to their reference collection since they diered in age, origin and, most relevantly, history. It was thus possible
to shed light upon the fossil shark species in Sicilian rocks and on their age and ndspot.
e draing of this paper was made possible thanks to Gaetano Giorgio Gemmellaro’s own research and, most of all,
to his son, Mariano.
Back in 1857, the former engaged in analysing fossil shes hitherto discovered in Sicily. Some of the nds – coming
from the provinces of Catania, Syracuse, Ragusa, Caltanissetta and Agrigento – were gathered by the scholar himself.
Mariano Gemmellaro’s work was rather more solid; between 1912 and 1913 he edited four monographs on Sicilian
Ichthyodontolites in which he studied fossil teeth of osseous and cartilagineous shes.
e material was collected by Mariano Gemmellaro himself, Giovanni Di Stefano and Giuseppe Checchia-Rispoli,
and is now entirely housed at Palermo’s Gemmellaro Geology Museum; while the remainder of the nds was already
stored at the museum.
In 1918, as director of the Geology Museum, Mariano Gemmellaro devoted himself to sorting and cataloguing those
many collections.
M. Gemmellaro’s untimely death in 1921 put an end to a research eld that was proving as stimulating just as rewarding.
His cataloguing work was not to be carried forward in the years ahead.
tHE cOLLEctiOns
C F T C (E – S)
e collection is made up of 6 teeth, belonging to 6 separate shark species from Patàra, between the towns of Trabia
and Termini Imerese (Palermo province).
Each nd has an illustration attached, itself drawn from Mariano Gemmellaro’s work (1912b). On the basis of macro-
foraminiferous (Nummulites) association, the author assigned the pieces to the Middle Eocene.
G T C (E – M - P – S)
e collection consists of 238 shark teeth and some vertebrae, all Sicilian in origin, and dating back to a no-better
specied “Mio-Pliocene“. 11 teeth are laid out in G. G. Gemmellaro’s 1857 article, 57 others are shown in M. Gemmellaro’s
papers (1912a; 1913a). Based on the palaeontological associations, M. Gemmellaro referred sharks to the Langhian.
e Ichthyodontolites unearthed in the Ragusa tar quarries may be related to the “Middle Miocene” (G
M., 1913a).
Q M F T L, S, ., (P – S)
e collection comprises 6 groups with 53 shark teeth, 22 of which illustrated by M. Gemmellaro (1913b); ey come
from Bagheria, Monte Pellegrino, Ficarazzi (all in the Palermo province) and Sciacca (Agrigento province) and were
found within the tufaceous limestone and clays of the Sicilian Plateau.
I V S S C
e collection consists of 46 allotments adding up to 300 shark teeth, together with large numbers of dental fragments,
one dermic plaque and a vertebra. Only 11 allotments come from Sicily.
FinDs PrOVEnAncE
e Ichthyodontolites fossiliferous layers may be traced back to a time wedged between the Middle Eocene (Lutetian)
and the Early Pleistocene (Sicilian). ey originate from a number of locations (g. 1).
Sicilian discoveries are henceforth listed and grouped together by identity or lithological analogy; they are set out in
chronological order according to the geological period they pertain. Each group is followed by a species up-to-date index
(C, 2002/2003).
112
eocene
T – T I (C/ P)
e selachimorph teeth from C/da Patàra refer to the colluvial layers of the Polizzi Fm. (Middle Eocene – Oligocene)
(A et al., 1988). Age: Middle Eocene (M. Gemmellaro,1912 b, Schmidt di Friedberg, 1964).
Located species:
Miocene
C, L, P, P A, B
e teeth from Corleone and the surroundings of Lercara and Prizzi arise from the Corleone Limestone Fm.
(R, 1966), consisting of glauconitic limestone intermeshed with green clayey-sandy levels, pertinent to the
Burdigalian –early Langhian stages (B & D P, 2001).
Such formation is a typical instance of open shelf deposit, deformed by Mio-Pliocenic compressive tectonics (N
& R, 1998).
e Corleone Limestone Fm. sits on levels of sandy clay, cancellophycus marl and lepidocycline limestone, dating to the
Middle Oligocene - Early Miocene (B & D P, 2001). Likewise, the nds from Palazzo Adriano and
Burgio belong to such context (C-R, 1911).
Located species:
Original determination Updated determination
Odontaspis macrota (Agassiz, 1843) Odontaspis macrota (Agassiz, 1843)
Odontaspis hopei (Agassiz, 1843) Odontaspis hopei (Agassiz, 1843)
Lamna obliqua (Agassiz, 1843) Odontaspis obliqua (Agassiz, 1843)
Oxyrhina desori (Agassiz, 1843) Isurus desori (Agassiz, 1843)
Carcharodon auriculatus (Blainv., 1818) Carcharodon megalodon (Agassiz, 1838)
Ginglymostoma priemi Gemm., 1912) Ginglymostoma priemi Gemm., 1912)
Original determination Updated determination
Odontaspis cuspidata (Agassiz, 1843) Odontaspis cuspidata (Agassiz, 1843)
Odontaspis contortidens (Agassiz, 1843) Carcharias sp.
Oxyrhina desori Agassiz, 1843 Isurus desori (Agassiz, 1843)
Oxyrhina hastalis Agassiz, 1843 Isurus hastalis (Agassiz, 1843)
Fig. 1 – Location of sites of origin of fossils
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R, S, M, M, V, M, G, V N
e nds from this district pertain the Ragusa Fm. (Upper Membro Irminio level, dating to the Aquitanian-Langhian
stages), made up of calcarenites with tar impregnations and white, grey, yellowish, well stratied marl-vaulted calcirudites.
Occasionally, the layers intermesh with marlic limestone (S F, 1964). e deposits are evidence of
a carbonate shelf neritic environment (B et al., 1987).
A number of researchers (G, 1857; M. G, 1913; D’E, 1920; B & L
R, 1962; R, 1962; S F, 1964) have brought up the issue of sh teeth within non-tar
and phosphate layers in and around Ragusa, Scicli and Modica. Despite the shortage of data, the pieces from Monterosso,
Vizzini, Militello, Val di Noto and Grammichele are to be conrmed within the Ragusa Formation group.
Located species:
P, S (F B)
M. Gemmellaro inferred (1913a) the high numbers of shark teeth from Pachino and Syracuse may have dated to
the Middle Eocene as well as to the Early and Middle Miocene. e former would be mirrored in the discontinuous
foraminiferous calcarenite Lutetian deposits; while the latter is represented by limestone layers rich in Aquitanian algae,
lepidocycline and bryozoids (C, 1963). Interestingly, D’E (1920) accounted for white-yellowish
Middle Miocene marlic limestone surfacing in this area. However, recent analysis (C et al., 1984) makes it likelier
the pieces are from the Membro dei Calcari calcarenite deposits in Syracuse, dating to the Middle Miocene.
Located species:
Syracuse:
Pachino:
Original determination Updated determination
Carcharodon megalodon Agassiz, 1838 Carcharodon megalodon Agassiz, 1838
Oxyrhina desori Agassiz, 1843 Isurus desori (Agassiz, 1843)
Original determination Updated determination
Odontaspis cuspidata (Agassiz, 1843) Odontaspis cuspidata (Agassiz, 1843)
Odontaspis contortidens (Agassiz, 1843) Carcharias sp.
Oxyrhina desori Agassiz, 1843 Isurus desori (Agassiz, 1843)
Oxyrhina hastalis Agassiz, 1843 Isurus hastalis (Agassiz, 1843)
Oxyrhina crassa Agassiz, 1843 Isurus sp.
Carcharodon auriculatus (Blainv., 1818) Carcharodon megalodon Agassiz, 1838
Carcharodon megalodon Agassiz, 1838 Carcharodon megalodon Agassiz, 1838
Hemipristis serra Agassiz, 1843 Hemipristis serra Agassiz, 1843
Original determination Updated determination
Odontaspis cuspidata (Agassiz, 1843) Carcharias sp.
Galeocerdo aduncus Agassiz, 1843 Galeocerdo aduncus Agassiz, 1843
Oxyrhina desori Agassiz, 1843 Isurus desori (Agassiz, 1843)
Oxyrhina hastalis Agassiz, 1843 Isurus hastalis (Agassiz, 1843)
Carcharodon auriculatus (Blainv., 1818) Carcharodon megalodon Agassiz, 1838
Carcharodon megalodon Agassiz, 1838 Carcharhinus sp.
Original determination Updated determination
Carcharodon auriculatus (Blainv., 1818) Carcharodon megalodon Agassiz, 1838
Hemipristis serra Agassiz, 1843 Hemipristis serra Agassiz, 1843
Carcharias egertoni (Agassiz, 1843) Carcharhinus egertoni (Agassiz, 1843)
114
P
In the immediate vicinity of Patti, Luigi Seguenza (1900) reported “Helvetian calcariferous greensand“ deposits
(Middle Miocene) containing selachimorph fossil teeth identical to those kept at the Museum in Palermo. In terms of
age and palaeontological contents, such lithology matches the Floresta Calcarenite Fm. (Late Burdigalian - Langhian),
surfacing in the Patti area.
Located species :
A, A, C
No better dating than a broad “Mio-Pliocene/Messinian” may be attached to the evidence from these sites. e piece
from the Adrano area (G, 1857) was gathered in a sulphureous marl context; the Cianciana nds were
extracted out of “…sulphureous areas, wihin the clay layers on top of the sulphur ones ”. e same applies to the Agrigento
discoveries. All may be dated to the Messinian.
Located species:
C G
No specication is given as to the one shark tooth from this location. It may well belong to a Carcharodon megalodon
individual, a common acquaintance amid the Miocenean soils of Sicily.
A
G. G. Gemmellaro (1857) studied two teeth from this town. On the strength of his own palaeontological deductions,
M. Gemmellaro later referred them to Miocenic levels.
Located species:
Pliocene
R, E
On the basis of M. Gemmellaro’s data (1913b), the acquisitions from Racalmuto and Enna were ascertained to the
Pliocene. No indication is given as to the assemblage lithology.
Located species:
Pleistocen
B, F, M P ( )
ese sites display the Late Pliocenic/Pleistocene carbonate silts making up the Palermo Plain. e deposits correspond
to the “limestone tufa“ (calcarenites) and clays quoted by M. Gemmellaro (1913b). ey have been narrowed down to the
Lower Pleistocene (Sicilian).
Original determination Updated determination
Odontaspis cuspidata (Agassiz, 1843) Carcharias sp.
Oxyrhina desori Agassiz, 1843 Isurus desori (Agassiz, 1843)
Oxyrhina hastalis Agassiz, 1843 Isurus hastalis (Agassiz, 1843)
Carcharodon auriculatus (Blainv., 1818) Carcharodon megalodon Agassiz, 1838
Original determination Updated determination
Oxyrhina spallanzani Bon., 1839 Isurus desori (Agassiz, 1843)
Oxyrhina hastalis Agassiz, 1843 Isurus hastalis (Agassiz, 1843)
Carcharodon rondeleti M. et H., 1841 Carcharodon megalodon Agassiz, 1838
Carcharodon megalodon Agassiz, 1838 Carcharodon megalodon Agassiz, 1838
Original determination Updated determination
Oxyrhina crassa Agassiz, 1843 Isurus sp.
Original determination Updated determination
Oxyrhina spallanzani Bon., 1839 Isurus oxyrhinchus Raf., 1809
Carcharodon rondeleti M. et H., 1841 Carcharodon carcharias (Linn., 1758)
Lamna sp. Isurus Hastalis (Agassiz, 1843)
Oxyrhina spallanzani Bon., 1839 Isurus desori (Agassiz, 1843)
115
Located species:
S
Two teeth come from this context. M. Gemmellaro (1913b) related the one from quaternary calcarenite deposits
around Sciacca to Odontaspis acutissima, while the second piece was assigned to Odontaspis cuspidata.
C
e only Sphirnidae remain in store comes from the Catania area. On the basis of the wealth of Pliocene and
Pleistocene, mainly bio-calcarenitic, deposits in this sub-region, we may place a pleistocenic tag on the nd. Such an
inference is strengthened by Sphirnidae stratigraphic patterns (B et al., 1987) (Cretaceous - Present).
Located species :
rEViEW
In the second half of the 19th century G. G. Gemmellaro embarked into the earliest attempt to classify the collections.
In the early 20th century, M. Gemmellaro reviewed such material and went onto catalogue the pieces gathered at a later
stage.
An untimely death (1921) prevented Mariano from updating his determinations according to the new scientic
theories that were to arise in the following decades.
Almost a century later, most of M. Gemmellaro’s cataloguing was found to be very consistent with only a handful of
updates in nomenclature needed; in other instances, the state of the pieces was so fragmentary as to prevent in-depth
analysis, thus calling for conrmation of old determinations.
As for the systematics of living Elasmobranchii, we hereby follow Compagno’s (1999) classication; while fossil
species were dealt with along the lines of Menesini (1969) and Landini’s (1977) papers. e Chondrichthy fauna is thus
displayed:
Original determination Updated determination
Sphirna prisca Agassiz, 1843 Sphirna prisca Agassiz, 1843
Original determination Updated determination
Odontaspis acutissima Ag., 1844 Odontaspis acutissima Ag., 1844
Oxyrhina spallanzani Bon., 1839 Isurus oxyrhinchus Raf., 180
Notidanus griseus Gmelin, 1788 Hexanchus griseus Hastalis (Bon., 1788)
Carcharodon rondeleti M. et H., 1841 Carcharodon carcharias (Linn., 1758)
C Chondrichthii
S Elasmobranchii
S-
Selachimorpha (Pleurotremata)
O Hexanchiformes Buen, 1926 G Carcharodon Agassiz, 1838
F Hexanchidae Gray, 1851 S Carcharodon carcharias (Linneo, 1758)
G Hexanchus Ranesque, 1810 Carcharodon megalodon Agassiz, 1838
S Hexanchus griseus (Bonaterre, 1788) O Carcharhiniformes Compagno, 1973
O Lamniformes Berg, 1958 F Carcharhinidae Jordan & Evermann, 1896
F Odontaspidae Müller et Henle, 1839 G Carcharhinus Blainville, 1816
G Carcharias Ranesque, 1810 S Carcharhinus egertoni (Agassiz, 1843)
S Carcharias sp. Carcharhinus sp.
G Odontaspis Agassiz, 1838 G Galeocerdo Müller et Henle, 1837
S Odontaspis cuspidata Agassiz, 1843 S Galeocerdo aduncus Agassiz, 1843
Odontaspis acutissima Agassiz, 1844 F Hemigaleidae Hasse, 1879
Odontaspis hopei (Agassiz, 1843) G Hemipristis Agassiz, 1843
Odontaspis macrota Agassiz, 1843 S Hemipristis serra Agassiz, 1843
116
: H R, 1810 (Fig. 2)
e Hexanchus genus consists of varieties known to date from as far back as the Jurassic. Currently only one species
survives, the Hexanchus griseus (Miocene–Recent).
is type of shark lives mainly in deep water, beyond 200 m and as far as 2000 m. While it primarily occupies tropical
and temperate waters, it may also be found in the Mediterranean (M, 1997).
In the collections, the Hexanchus genus only appears in the form of a single tooth, which M. Gemmellaro (1913)
ascribed to the living Hexanchus griseus.
: C R, 1810 - O A, 1838 (Figg. 3, 4, 5)
Odontaspids, known of since the Cretaceous, usually inhabit warm sub-tropical water and tend to lodge in sandy,
neritic sea bottoms (C, 1972).
e collection holds around 100 teeth, almost all rootless and assigned to various species within the Odontaspis genus.
In the absence of roots, themselves a key to the species taxa, it is likely that most determinations are incorrect and that most
of the pieces are to be related to Carcharias sp. Among the ascertained Odontaspis specimen, there are both Odontaspis
cuspidata and Odontaspis acutissima.
Odontaspis acutissima is known through the Belgian, North Italian and West German Oligocene. It was very widespread
both in the Mediterranean basin and in the Atlantic, and was still fairly abundant there up to the end of the Pliocene
(M, 1971). In Sicily, this species appears in the Early Pleistocene.
Odontaspis cuspidata rst springs up in the Late Palaeocene; it develops widely both in the Mediterranean and the
Atlantic; only to face extinction in the Pliocene.
Lastly, it is worthy to note the teeth M. Gemmellaro determined as Odontaspis macrota, Odontaspis hopei and
Odontaspis obliqua, actually all belong to the Odontaspis taurus obliqua subspecies (C, 1972).
G : I R, 1810 (Figg. 6, 7, 8)
e Isurus genus, originating in the Early Cretaceous, is currently represented by two species; one of the them – Isurus
oxyrhinchus – also dwelling the Mediterranean. e Isurus oxyrhinchus lives in all warm regions, and partly in temperate
ones (Pliocene–Recent); it has a long slim gure and large dimensions.
is shark occupies coastal and ocean waters alike, at depths never greater than 150 m. It may be encountered all
around the globe so long as the water temperature is above 16° C (M, 1997). Isurus is widespread in neogenic
soils, particularly within neritic bottoms, and is almost always associated with Carcharodon (s.l.) (L, 1977).
is genus is represented in the collection through 150 teeth and is the most commonly featured shark in the Sicilian
catchment area. ree species have been singled out: Isurus hastalis (61 teeth), Isurus desori (58 teeth), Isurus oxyrhinchus
(6 teeth).
e attribute enabling to distinguish shark species is the root complex; however, given its absence in the vast majority
of instances, more oen than not classication is problematic.
Isurus hastalis (Oligocene – Pliocene) is the second best known species aer Carcharodon megalodon, both in terms
of its distribution (Europe, Africa, North- and South-America, New Zealand, Japan) as well as in the number of fossil
resources. In analysing Isurus hastalis dental morphology and features – particularly the miocenic specimens – many
researchers have found this species to be bigger built than today’s equivalent. Caretto (1972) has it that Isurus hastalis’
larger size may be due to:
1. Larger dimensions of Cenozoic individuals.
2. Favourable climatic conditions as to allow a greater development of the single individuals.
Isurus desori (Eocene – Pliocene) displays larger teeth than those of Isurus hastalis, yet they may not be easily
discriminated from one another owing to the lack of root complexes. is species is known to have existed since the
Eocene, it spread out widely during the Miocene (Europe, North-Africa, South-America), and nally shrunk solely to
Argentina in the Pliocene.
Odontaspis obliqua (Agassiz, 1843) F Sphirnidae Gill, 1872
F Lamnidae Müller et Henle, 1838 G Sphirna Ranesque, 1810
G Isurus Ranesque, 1810 S Sphirna prisca Agassiz, 1843
S Surus oxyrhinchus Ranesque, 1809 O Orectolobiformes Applegate, 1972
Isurus hastalis (Agassiz, 1843) F Ginglymostomatidae Gill, 1862
Isurus desori (Agassiz, 1843) G Ginglymostoma Müller et Henle, 1837
Isurus sp. S Ginglymostoma priemi Gemmellaro, 1912
117
Fig. 2 – Hexanchus griseus. Pleistocene, Ficarazzi (PA). Coll. “Fossili quaternari dei tu calcarei e delle sabbie marine”.
Fig. 3 – Carcharias sp. Miocene, Ragusa. Coll. “Gemmellaro Terziario”.
Fig. 4 – Odontaspis acutissima. Pleistocene, Monte Pellegrino (PA). Coll. “Fossili quaternari dei tu calcarei e delle sabbie marine”.
Fig. 5 – Odontaspis macrota. Eocene, Termini Imerese (PA). Coll. “Crostacei e denti di pesce”.
Fig. 6 – Isurus hastalis. Miocene, Pachino (SR). Coll. “Gemmellaro Terziario”.
Fig. 7 – Isurus desori. Miocene, Ragusa. Coll. “Gemmellaro Terziario”.
Fig. 8 – Isurus oxyrhinchus. Pleistocene, Ficarazzi (PA). Coll. “Fossili quaternari dei tu calcarei e delle sabbie marine”.
Fig. 9 – Carcharodon megalodon. Miocene, Siracusa. Coll. “Gemmellaro Terziario”.
Fig. 10 – Carcharodon carcharias. Pleistocene, Palermo. Coll. “Fossili quaternari dei tu calcarei e delle sabbie marine”.
Fig. 11 – Galeocerdo aduncus. Miocene, Ragusa. Coll. “Gemmellaro Terziario”.
Fig. 12 – Hemipristis serra. Miocene, Burgio (AG). Coll. “Gemmellaro Terziario”.
Fig. 13 – Ginglymostoma priemi. Eocene, Termini Imerese. Coll. “Crostacei e denti di pesce”.
118
G : C A, 1838 (Figg. 9, 10)
Only one, large, cosmopolitan, migratory species, the Carcharodon carcharias (Linneo, 1758), belongs to this genus.
It may be found in all temperate-warm waters (C, 1972).
Carcharodon carcharias (Pliocene–Present) is stoutly built, elongated and broadly mouthed, with triangular saw-like
teeth (M, 1997). It inhabits all areas, from temperate waters to the oceans (Mediterranean, Pacic, Atlantic), at
depths between the surface and 250 m; it is a great sea predator and is oen to be spotted along the coastline.
Twelve Carcharodon carcharias teeth are on stock, together with over 80 others belonging to the extinct species
Carcharodon megalodon, Agassiz 1838. Despite researchers now tend to discard philogenetic links, the two do share
dental setting and general shape: so much so that they may only be distinguished through average size mean (N &
P, 1988). In fact Charcarodon megalodon teeth show an isosceles triangle shape, a saw-like pattern along the edge
and, most of all, remarkable height of up to 15 cm.
e species gradually expanded between the Eocene and the Oligocene, it dilated signicantly in the Miocene, and
disappeared at the start of the Pliocene. As to geography, Charcarodon megalodon is one of the most easily encountered shark
forms, though altogether tied up to Odontaspis and Isurus (especially Isurus hastalis). During the Miocene – particularly
so in the earlier stages of this epoch – the population swell remarkably and expanded further aeld (C, 1972).
G: C B, 1816
Current Carcharhiniformes include sharks of varying dimension, migratory as well as coast-settlers, lodging in all
marine contexts, neritic and deep-water (C, 1972).
Four teeth are held in stock, three of which belong to the extinct Carcharhinus egertoni. is shark is known of since
the Miocene, both in the Mediterranean and in the Atlantic. In the Pliocene, it conned itself to the Mediterranean basin
alone (M, 1971).
G: G M H, 1837 (Fig. 11)
At present, only Galeocerdo cuvieri (Péron & Lesueur, 1822) falls within this genus. is large shark lives on the coast,
between the surface and 140 m in the temperate – tropical waters of the Atlantic and of the Indo-Pacic region (M,
1997). Only one tooth is stored at the Museum, which M. Gemmellaro (1913) referred to the extinct Galeocerdo aduncus,
Agassiz 1843. Caretto (1972) observes that these teeth are smaller than in today’s species and represent slightly smaller,
though comparable, individuals.
Galeocerdo aduncus rises in the Eocene through and ripens in the Miocene, with a greater diusion in the latter
epoch.
G : H A, 1843 (Fig. 12)
Hemipristis serra (Oligocene – Miocene) appears in the collection with 9 entries, very similar to the present morphology,
even if generally smaller sized (C, 1972).
Hemipristis serra is an oen-found, typically neogenic species, bound to other forms like Lamniformes and
Odontaspids; moreover, Menesini (1969) suggests that during the Miocene this may well have been regarded as a globally
diused species.
G: S R, 1810
Only a single rootless tooth is kept of the extinct Sphirna prisca. Owing to the lack of the root complex, this specimen
is likely to be related to Carcharhiniformes rather than to Sphirna for many crown characteristics are common to the
two genera. Sphirna prisca appears in the Oligocene and it is a common acquaintance in Miocenic Mediterranean and
Atlantic layers, only to become rarer in the Miocene.
At present, the Sphirna genus appears in all temperate-tropical seas (including the Mediterranean); it lives on
and o the coast at depths of up to approximately 300 m.
G: G M H, 1837 (Fig. 13)
e Ginglymostoma genus was rst assigned to Sicily by M. Gemmellaro’s (1912b) discovery of a small, incomplete
tooth. He recorded the nd as a Ginglymostoma priemi, thus establishing a new species. Ginglymostoma specimens are rare
and for this reason the original determination is being maintained.
PALAEOcLiMAtic AnD PALAEOEcOLOGicAL MODELLinG
Palaeoenvironmental rendering may be attempted on the grounds of customs observed on individuals from living
species; extinct species are to be modelled on the account of their closest (living) successors.
On commencement, we shall like to stress that only three genera of all (Isurus, Carcharodon e Odontaspis) feature
copiously, being derived from middle-large sized specimens. at may be owed to:
119
1. e greater endurance of large fragments to diagenetic processes;
2. A tendency in the past to gather and preserve larger nds.
Such occurrence would not therefore rule out the presence of equal numbers of smaller built species and/or younger
individuals from those very large species: it may also account for their reduced rate of preservation.
ere are not biostratigraphic data and no detailed sedimentological and palaeontological inference may be drawn.
e evidence represents too narrow a species sample to be of any signicance to palaeoenvironmental modelling.
Likewise, several species are ill represented, thus making environmental analysis far from feasible.
All of the above bears the impediments involved in rendering palaeoclimate and ecology. However, one useful piece
of data that should be not discounted is the common lithological horizon of most nds: calcarenites and bio-calcarenites,
indicating shallow waters (i.e. a neritic region). e presence of some open-water shark allow us to consider these
palaeoenviroment such as an ideal hunting areas.
A greater deal of attention should be paid to the Early-Middle Miocene layers. ese are the deposits yielding the
greater number of nds and the broader faunal variety. e specimens there in unearthed exemplify warm sub-tropical
forms.
Coastal and open-water sharks co-existed side-by-side in the record. Parallel studies on other Italian regions (Apulia,
Piedmont and the Central Apennine range) have thrown up the very same faunal associations within similar litho-types.
Such relations collate with the existence of warm, closed, shallow seas in the Early-Middle Miocene. Moreover, faunal
anities may easily be framed within Italy’s contemporary palaeogeographic situation, whereby a short stretch of sea
would have been linking the North West of the peninsula to Sicily, via the Apennine range.
ree of the four species from the Early Pleistocene deposits around Palermo (Bagheria, Ficarazzi, Monte Pellegrino)
and Sciacca are still living. is may point to a faunal association closer to that currently present in the Mediterranean; we
may also infer its likely origin to be traced back to a warm-temperate, neritic environment.
Rounding up, the fossil evidence may assist to support – though along very broad lines – earlier theories based upon
geochemical (δ18 O) and palaeontological in-depth examination (M et al., 1998; J et al., 2003).
cOncLusiOns
Current analysis may allow us to isolate:
– Several instance displaying little or no data alongside fully referenced (provenance, location, dating) nds by
Gaetano Giorgio Gemmellaro and Mariano Gemmellaro,
– e bulk of the evidence may be dated to between the Middle Eocene (Lutetian) and the Lower Pleistocene
(Sicilian), with larger concentrations during the Early and Middle Miocene (Aquitanian - Langhian).
– Fossilifereous resources are primarily located in the Central-Western and South-Eastern corners of the Island.
– e majority of nds pertains to a common lithological horizon (limestone and greenstone platform contexts).
– Large sized species seem to be over-represented, particularly so in the instance of Isurus, Carcharodon and
Odontaspis remains.
– e review brings truth to the widespread presence of sharks in the Italian Cenozoic, especially in reference to
the Miocene, on the grounds of close faunal anities in Sicily and other regions (Apulia, Piedmont and Tuscany).
– In terms of palaeoclimate and palaeoecology, the bulk of faunal assemblages – though not all from contemporary
contexts – indicates their origin to be within neritic habitats, with warmer waters than present and rich prey. Sound
evidence of such situation lies in the abundance of osseous shes both in the collections and in the limestone layers
associated to shark teeth.
Lastly, faunal assemblages may then be taken as to conrm the palaeoenvironmental framework drawn upon more
consistent data (Foraminifera, Molluscs and Echinoids).
AcKnOWLEDGMEnts
e authors wish to thank Prof. Walter Landini and Prof. Franco Cigala–Fulgosi for their assistance in carrying out
the review of the manuscript.
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