Historical Biology

An International Journal of Paleobiology

ISSN: (Print) (Online) Journal homepage: https://www.tandfonline.com/loi/ghbi20

An overview of the fossil turtles from Sardinia

(Italy)

Daniel Zoboli, Georgios L. Georgalis, Marisa Arca, Caterinella Tuveri,

Salvatore Carboni, Luciano Lecca, Gian Luigi Pillola, Lorenzo Rook, Mauro
Villani, Francesco Chesi & Massimo Delfino

To cite this article: Daniel Zoboli, Georgios L. Georgalis, Marisa Arca, Caterinella Tuveri,
Salvatore Carboni, Luciano Lecca, Gian Luigi Pillola, Lorenzo Rook, Mauro Villani, Francesco
Chesi & Massimo Delfino (2022): An overview of the fossil turtles from Sardinia (Italy), Historical
Biology, DOI: 10.1080/08912963.2022.2098488

To link to this article: https://doi.org/10.1080/08912963.2022.2098488

Published online: 29 Jul 2022.

Submit your article to this journal

View related articles

View Crossmark data

Full Terms & Conditions of access and use can be found at

https://www.tandfonline.com/action/journalInformation?journalCode=ghbi20
HISTORICAL BIOLOGY
https://doi.org/10.1080/08912963.2022.2098488

An overview of the fossil turtles from Sardinia (Italy)

Daniel Zoboli a, Georgios L. Georgalis b,c, Marisa Arcad, Caterinella Tuverid, Salvatore Carbonia, Luciano Leccaa,
Gian Luigi Pillolaa, Lorenzo Rook e, Mauro Villanif, Francesco Chesie and Massimo Delfino g,h
a
Dipartimento di Scienze Chimiche e Geologiche, Università di Cagliari, Cittadella Universitaria, Monserrato, Italy; bInstitute of Systematics and Evolution
of Animals, Polish Academy of Sciences, Kraków, Poland; cPalaeontological Institute and Museum, University of Zurich, Zurich, Switzerland;
d
Soprintendenza Archeologia, Belle Arti e Paesaggio per le Province di Sassari e Nuoro, Nuoro, Italy; eDipartimento di Scienze della Terra, Paleo[Fab]Lab,
Università di Firenze, Florence, Italy; fMuseo del carbone – Centro italiano della cultura del carbone (CICC), Grande Miniera di Serbariu, Carbonia, Italy;
g
Dipartimento di Scienze della Terra, Università di Torino, Turin, Italy; hCarrer de les Columnes s/n, Campus de la UAB, Institut Català de Paleontologia
Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, Barcelona, Spain

ABSTRACT ARTICLE HISTORY

We review in detail the published fossil record of turtles from Sardinia and, in addition, we document Received 22 May 2022
previously undescribed specimens for the first time. Among these undescribed specimens, is the oldest Accepted 3 July 2022
occurrence of Testudo hermanni on the island, from the Early Pleistocene of Monte Tuttavista. The turtle fossil KEYWORDS
record in Sardinia goes back to the Eocene and comprises 18 different taxa, pertaining to 6 lineages: Testudines; taxonomy;
Podocnemididae, Cheloniidae, Emydidae, Geoemydidae, Testudinidae, and Trionychidae. Remarkable is palaeobiogeography;
the occurrence of Eocene pleurodires, whose presence is in agreement with the Oligo-Miocene rifting of Cenozoic; Italy
the Corso-Sardinian block. Interestingly, the fossil record provides evidence for the presence in the island of
both Testudo hermanni and Emys orbicularis during the Pleistocene although according to molecular data
the extant populations of these two taxa were introduced in recent times. Finally, a large ungual phalanx
from the Middle–Late Pleistocene Monte San Giovanni bone breccia testifies the occurrence of a giant
tortoise in the Quaternary terrestrial ecosystem of Sardinia.

Introduction
Palaeogeographic context
The Sardinia-Corsica Massif represents a large segment of the
Europe foreland and Alpine orogenic system originally located
in continuity with the southern European continental plate (see
among others Vardabasso 1962; Cherchi 1974; Speranza et al.
2002; Gattacceca et al. 2007; Oudet et al. 2010; Advokaat et al.
2014). From the Oligocene onwards, this area started to sepa­
rate from the mainland with a volcanogenic rifting stage fol­
lowed by a counterclockwise rotation (Aquitanian–early
Burdigalian) that concluded during the Middle Miocene
(Langhian). Different insular vertebrate faunas are reported
during the Neogene, among which the oldest known from the
whole Mediterranean Sea (De Bruijn and Rümke 1974; Van der
Made 2008; Mennecart et al. 2017, 2019). Other insular verte­
brate faunas are documented in the Late Miocene (Abbazzi
et al. 2008a; Casanovas-Vilar et al. 2011) and in the Plio-
Pleistocene (Van der Made 1999; Abbazzi et al. 2004, 2008b;
Zoboli and Pillola 2016a; Palombo 2018).
An overview of the Cenozoic record of Sardinian reptiles
The Cenozoic reptiles of Sardinia are represented by the higher taxa
that currently inhabit the Northern Hemisphere: Crocodylia,
Testudines, and Squamata (i.e., lizards – including worm lizards –
and snakes).
Crocodylian remains have been reported from the early Eocene
to the Late Miocene and are generally represented by isolated teeth
referred to Crocodylia indet. (Del Vecchio 1921; Abbazzi et al.
2008b; Zoboli et al. 2019), and rare cranial and postcranial elements
(Gennari 1868; Capellini 1890a,b; Spano 1985). The most diagnos­
tic remains consist of an incomplete cranium and few postcranial
elements from a single specimen collected in the Miocene marine
succession of Cagliari (Gennari 1868), which allowed the descrip­
tion of the new tomistomine crocodylian ‘Tomistoma’ calaritanum
Capellini, 1890b whose specific validity has been recently confirmed
by Nicholl et al. (2021).
Lizards have been reported from the Early Miocene to the
Holocene (Venczel and Sanchiz 2006; Delfino et al. 2011).
A form similar to the anguid lizard Ophisaurus fejfari
Klembara, 1979 was reported from the Early Miocene lacustrine
deposit of Oschiri (northern Sardinia) (Ophisaurus cf. O. fejfari
of Venczel and Sanchiz 2006), whereas Ophisaurus spinari
Klembara, 1979 is present in the Late Pliocene of Capo
Mannu D1 Local Fauna (central western Sardinia) (Delfino
et al. 2011; Klembara and Rummel 2018). Other lizard taxa
were reported from several Pleistocene fissure fillings at Monte
Tuttavista (central eastern Sardinia): Agama s.l. sp., Timon sp.,
Podarcis sp., Lacerta sp., and Gekkonidae indet. (Abbazzi et al.
2004; Delfino et al. 2008; Tschopp et al. 2018). Very few
remains of skinks are known from the Early Miocene of
Oschiri to the Holocene of few localities (Delfino 2002;
Venczel and Sanchiz 2006). Among worm lizards, Blanus gra­
cilis (Roček, 1984) was reported from the Early Miocene of
Oschiri (Venczel and Sanchiz 2006), and the remains of unde­
termined amphisbaenians were collected in the Late Pliocene of
Capo Mannu D1 Local Fauna (Delfino et al. 2011) and in the
Pleistocene fissure fillings at Monte Tuttavista (Abbazzi et al.
2004).

CONTACT Daniel Zoboli danielc.zoboli@unica.it Dipartimento di Scienze Chimiche e Geologiche, Università di Cagliari, Cittadella Universitaria SS-554, Monserrato
09042, Italy
© 2022 Informa UK Limited, trading as Taylor & Francis Group

Published online 29 Jul 2022

2 D. ZOBOLI ET AL.
Miocene snakes are represented by a few taxa from Oschiri:
Eoanilius oligocenicus Szyndlar, 1994, Vipera gr. V. aspis
(Linnaeus, 1758), and Natricinae indet. (Venczel and Sanchiz
2006), however, none of them have so far been figured.
Recently, the type material of the purported constrictor snake
Palaeopython sardus Portis, 1901a from the Miocene of Bosa
(central western Sardinia) (Portis 1901a) was referred to an
undetermined acanthomorph fish (Delfino et al. 2014). In any
case, constrictor snakes (sensu Georgalis and Smith 2020) were
present in Sardinia during the Late Pliocene as testified by the
occurrence of Eryx cf. E. jaculus (Linnaeus, 1758) from Capo
Mannu D1 Local Fauna (Delfino et al. 2011). In addition,
Vipera sp., Natrix sp., and ‘Colubrinae’ indet. were reported at
Capo Mannu D1 Local Fauna (Delfino et al. 2011). Remains of
Pleistocene snakes, represented by isolated vertebrae, were also
collected in several Pleistocene cave deposits and fissure fillings
(Kotsakis 1980; Abbazzi et al. 2004). In the fissure fillings at
Monte Tuttavista were reported Natrix sp., Vipera sp. (Abbazzi
et al. 2004), and the Sardinian endemic colubroid Sardophis
elaphoides Georgalis and Delfino(Georgalis et al., 2019).
In this study, all published fossil Testudines of Sardinia are listed
and reviewed, with the addition of unpublished material (Figure 1).
Materials and methods/anatomical nomenclature
The anatomical nomenclature follows Zangerl (1969) and Gaffney
(1996), Hervet (2000), Joyce and Bell (2004), Joyce (2007), Vitek
and Joyce (2015), and Vlachos and Rabi (2018).
Nomenclature of turtle clades follows Joyce et al. (2021).
Taxonomy and nomenclature of extant turtle genera and species
follow Rhodin et al. (2021).
Anatomical structure abbreviations: Carapace and plastron
plates: C, costals; N, neurals; P, peripherals; PY, pygal; EPL, epiplas­
tra; ENTO, entoplastron; HYO, hyoplastron. Carapace and plastron
scutes: ce, cervical; ve, vertebrals; ple, pleurals; ma, marginals; s,
supracaudals; gu, gulars; hu, humerals.
Institutional abbreviations: MDLCA, Museo Sardo di
Geologia e Paleontologia ‘Domenico Lovisato’, University of
Cagliari, Italy; UC, University of Cagliari; CL, Carboni-Lecca
Collection of MDLCA; MARTEL, Museo dei Palaeoambienti
Sulcitani – ‘E.A. Martel’ of Carbonia, Italy; MPSMC, Museo
Civico Paleontologico e Speleologico ‘E.A. Martel’ of Carbonia
(currently MARTEL); MSNM, Museo Civico di Storia
Naturale of Milan, Italy; FS, field inventory register of
Fiume Santo material; MAN, Museo Archeologico Nazionale
‘Giovanni Asproni’ of Nuoro (Soprintendenza Archeologia,
Belle Arti e Paesaggio per le province di Sassari e Nuoro),
Nuoro, Italy; NMB, Naturhistorisches Museum Basel,
Switzerland; Ty., C.I.F. Major's Collection of NMB.
The fossil Testudines of Sardinia
Paleogene
Locality
Monte Sinni coal mine (Nuraxi Figus, Municipality of Gonnesa,
Iglesiente subregion, southwestern Sardinia).
Geological setting and age
Lignitifero Formation (early Eocene [late Ypresian–early
Lutetian]; Cita et al. 2007). The Sulcis Basin, in the southwes­
tern Sardinia, is characterised by a Paleogene succession (early
Ypresian–Rupelian). These deposits discordantly lie on the
Variscan basement and at the top are generally covered by an
Oligo-Miocene volcanic succession. Three formations are dis­
tinguished from bottom to top: the Miliolitico Fm. (early
Eocene [early Ypresian]), the Lignitifero Fm. (early–middle
Eocene [late Ypresian–early Lutetian]) and the Cixerri Fm.
(middle Eocene–early Oligocene [Lutetian–Rupelian]).
The Lignitifero Fm. (Cita et al. 2007) is generally badly exposed and
the most representative outcrops are located between Carbonia and the
village of Gonnesa. However, the succession is well known due to the
numerous mining surveys since the beginning of the 20th century. It
consists of rhythmic alternations of decimetre or metric layers of marls,
marly limestones, bituminous limestones, carbonaceous clays, lignite
layers, sandstones and microconglomerates. Carbonates are more
common in the lower part of the succession where marly limestones
rich in miliolids testify numerous and repeated marine ingressions.
Palaeosols are also locally associated with the first layers of lignite.
Upwards, lake-marsh facies with ostracods, characee, and molluscs
become more frequent. The lignite layers are less numerous but
thicker, and alluvial intercalations, represented by sandstones and
conglomerates, become more common (Cocozza et al. 1986). The
maximum thickness of the lignite layers usually does not exceed
20 m, but a thickness of more than 100 m is reported in the western
surveys, probably due to reverse faults and tilted strata (Fanni et al.
1982).
The palaeontological assemblage consists of macroflora (e.g.,
Sabal sp., Juglans sp.) and microflora (Myricaceae, Palmae,
Pteridophyta, Cupressaceae, Betulaceae, Fagaceae, Corylaceae, and
Juglandaceae) (Cita et al. 2007). The pollen association and the
presence of the Characeae Nitellopsis thaleri thaleri indicate a late
Ypresian–early Lutetian age for the Lignitifero Formation (Agus
and Pecorini 1978; Barbieri and Cherchi 1980). Fishes (cf. Alestes
sp.), reptiles (Pan-Trionychidae indet., see below), and mammals
(Amphiperatherium sp. and ‘Lophiodon’ sardus Bosco, 1902) are
reported (Bosco 1902; Cappetta and Thaler 1974; Kotsakis 1985;
Kotsakis et al. 1997).
The Monte Sinni coal mine was the only coal mine still active in
Italy in 2016, and the mining activity stopped definitively in 2019.
During the mining activity in the 90ʹs, the miner Mr Pasquale
Scotto found some turtle remains in the Seruci slope ‘Chilotta
Level’ at the depth of 350 m. The fossils were donated to
MARTEL, where they are currently stored.
Pan-Trionychidae indet.
Figure. 2
Material
MARTEL 0059, two fragments of costals from a single individual
and a possible incomplete humerus (Figure 2).
Description and remarks
Both costals are much longer than wide. They bear a distinctive
sculpturing pattern in dorsal view, which consists of dense and
distinct ridges. That ornamentation is more prominent in the
central part of the costal and slightly fades out towards its margins.
A further isolated skeletal element could represent a Pan-
Trionychidae humerus but due to its preservational status it cannot
be referred to this taxon with confidence.
The presence of a sculpturing pattern along with the absence of
shell scutes allow a referral of the specimen to Pan-Trionychidae
(Vitek and Joyce 2015; Georgalis and Joyce 2017).

Figure 1. Sardinian Cenozoic Testudines-bearing localities (the municipality is indicated
in brackets when it does not correspond to the name of the locality, the Holocene
specimens are from archaeological contexts from Neolithic to Middle Ages): 1, Monte
Sinni Mine (Gonnesa); 2, Bacu Abis (Carbonia); 3, Murecci (Gonnesa); 4, Medau is Fenus
(Carbonia); 5, San Michele (Laerru); 6, Nulvi; 7, Noragugume; 8, Bosa; 9, Fiume Santo
(Porto Torres); 10, Sassari; 11, Is Mirrionis-Piazza d'Armi (Cagliari); 12, Sant'Avendrace
(Cagliari); 13, Nuraghe Su Casteddu (Dorgali); 14, Capo Mannu D4 Local Fauna (San Vero
Milis); 15, fissure filling VII 2 – ‘mustelide’ and fissure filling VII – ‘giacimento 2’ of Monte
Tuttavista (Orosei); 16, Punta del Quadro (Alghero); 17, Monte San Giovanni (Iglesias/
Gonnesa); 18, San Giovanni di Sinis (Cabras); 19) Grotte di Is Zuddas (Santadi); 20, Grotta
di Monte Meana (Santadi); 21, Grotta Corbeddu (Oliena); 22, Grotta Su Guanu (Oliena);
23, Grotta Filiestru (Mara); 24, Monte Sant'Antonio (Siligo); 25, Turris Libisonis (Porto
Torres); 26, S' Imbalconadu (Olbia); 27, Santa Filitica (Sorso); 28, Santissima Trinità di
Saccargia (Codrongianos).
HISTORICAL BIOLOGY 3
Locality
Bacu Abis (Municipality of Carbonia, Sulcis subregion).
Geological setting and age
Lignitifero Formation (early Eocene [late Ypresian–early Lutetian];
Cita et al. 2007, see above). Kotsakis (1985) reported turtle remains
on a sandy level interbedded in a lignite sequence outcropping in
the Bacu Abis ancient mine. In the same horizon of this locality,
a few teeth of an herpetotheriid marsupial (Amphiperatherium sp.)
were discovered (Kotsakis et al. 2008a).
Pan-Trionychidae indet.
Material
Repository unknown, fragments of a carapace.
Description and remarks
Kotsakis (1985) did not figure the material from Bacu Abis
but reported that its carapace ornamentation is very similar to
Trionyx michauxi Broin, 1977, from the early Eocene of
northern France, a taxon that is currently considered
a junior synonym of ‘Trionyx’ silvestris Walker and Moody,
1974, from the early Eocene of England (see Georgalis and
Joyce 2017). In any case, the ornamentation pattern is
a highly variable feature within pan-trionychids (Gardner
and Russell 1994; Georgalis and Joyce 2017). As such, con­
sidering also the absence of any published figure, we follow
Georgalis and Joyce (2017), who treated this Sardinian mate­
rial as an indeterminate pan-trionychid. The whereabouts of
this material are currently not known (it is not present in the
collections of the Museo Universitario di Scienze della Terra,
Sapienza Università di Roma; R. Sardella pers. comm. to MD
22/07/2021).
Locality
Medau is Fenus, near Flumentepido (Municipality of Carbonia).
Geological setting and age
Cixerri Formation (probably middle–late? Eocene; Georgalis et al.
2020b). The Cixerri Fm. is a terrigenous formation represented by
siltites, sandstones, conglomerates, subordinated lacustrine lime­
stones and rare intercalations of lignitiferous clays (Pecorini and
Pomesano Cherchi 1969; Costamagna and Schäfer 2013). The facies
indicate a continental depositional context with a prevalence of
fluvial deposits and subordinate lacustrine levels (Pecorini and
Pomesano Cherchi 1969). The formation represents the molassic
post-Pyrenean phase sedimentation in Sardinia and deposed in the
middle to lower, distal part of a foreland alluvial plain fed by Iberian
relieves (Cherchi 1979; Costamagna and Schäfer 2013). In the
Sulcis–Iglesiente area, the succession rests conformably to uncon­
formably over the late Ypresian–early Lutetian Lignitifero Fm. or it
is placed unconformably over the Variscan metamorphic basement.
Upwards, it passes sharply through an unconformity to the cal­
calkaline volcanites of the Oligo-Miocene cycle (about
22.8 ± 1.3 Ma in the Sulcis area; Lecca et al. 1997), and it is intruded
by volcanic rocks dated at 29.30 ± 1.2 to 26 Ma (Funedda et al.
2009).
The palaeontological record is rather poor and consists of
rare plant remains, freshwater gastropods (Stagnicola cf.
S. orelongo), and undetermined bone fragments collected in
4 D. ZOBOLI ET AL.

Figure 2. Pan-Trionychidae indet. (MARTEL 0059), Monte Sinni coal mine (Gonnesa), late Ypresian–early Lutetian (Lignitifero Fm.). a-b, costals in external view.

lacustrine facies (Taricco 1924; Maxia 1959). The age of the Eocenochelus cf. E. eremberti
Cixerri Fm. is bracketed between the early Lutetian (which is
the age at the top of the Lignitifero Fm.) and the first local
occurrence of intruded mid-late Rupelian volcanic products. Figure. 3a
According to these data and the continental turtle biostrati­
graphy, the remains from the Cixerri Fm. have been referred
to the middle–late? Eocene (Georgalis et al. 2020b), but a late Material
Eocene age cannot be ruled out. The turtle specimen from MDLCA 14006, an almost complete carapace and plastron, missing
Medau is Fenus originates from a sand quarry opened near only tiny fragments of the anteriormost and posteriormost portions
the village of Flumentepido. of the carapace (Figure 3a).
 HISTORICAL BIOLOGY 5

Figure 3. a, Eocenochelus cf. E. eremberti (MDCLA 14006), Medau is Fenus (Carbonia), middle–late? Eocene (Cixerri Fm.). Almost complete carapace and plastron in dorsal
(a1), ventral (a2), and left lateral (a3) views. b, cf. Eocenochelus sp. (MDCLA 3018), Murecci (Gonnesa), middle-late? Eocene (Cixerri Fm.), incomplete carapace and plastron in
dorsal (b1) and ventral (b2) views. For an interpretative drawing see Georgalis et al. (2020b).
6 D. ZOBOLI ET AL.
Description and remarks
The specimen, an almost complete shell, has been recently
described in detail by Georgalis et al. (2020b). The carapace
misses only tiny fragments of the anteriormost and posterior­
most portions, especially at the posterior right portion, near the
marginals, as well as near the level of the first pleurals. Certain
portions of the carapace are slightly deformed and three
breakages are apparent in the carapace. Georgalis et al.
(2020b) referred the specimen from Medau is Fenus to erym­
nochelyine pleurodires; more in particular, they highlighted
strong resemblance with Eocenochelus eremberti (Broin, 1977)
from the middle Eocene of France, Belgium, and Spain, on the
basis of the following character combination: sinuous lateral
margins of the vertebral scutes; relatively long epiplastral sym­
physis, less than half the entoplastral length; relatively wide
posterior plastral lobe; subrounded lateral margins of the pos­
terior plastral lobe; relatively wide anal notch, its length being
equivalent to half its width or less (Georgalis et al. 2020b).
Accordingly, the specimen was referred to as Eocenochelus cf.
E. eremberti.
The occurrence of Eocenochelus in the middle Eocene of
Sardinia represents another shared faunal element between the
Island and continental Western Europe during the Paleogene and
provides interesting biogeographic implications (Georgalis et al.
2020b).
Locality
Murecci (Municipality of Gonnesa).
Geological setting and age
Cixerri Formation (probably middle–late? Eocene; Georgalis et al.
2020b, see above). The turtle remains from this locality originate
from a natural outcrop of the Cixerri Fm.
Figure 4. Pan-Trionychidae indet. (MDLCA 14116), San Michele (Laerru), early–
middle Burdigalian (Perfugas Fm.). Undetermined bone fragments and probable
costals.
cf. Eocenochelus sp.
Figure. 3b
Material
MDLCA 3018, an incomplete shell (Figure 3b).
Description and remarks
The specimen, recently described by Georgalis et al. (2020b), con­
sists of a partial shell, lacking the anterior portions of both carapace
and plastron. The preserved portion of the shell is filled with
a microconglomerate matrix and is locally heavily fractured and
incomplete. The surface is partly covered by a hard concretion.
Most of the boundaries of shell skeletal elements are not visible,
though their presence can be supposed based on the general shape
of the shell.
The main characters available in the shell from Murecci are
compatible with those observed in the specimen from Medau is
Fenus. The differences observed between them are recognised as
compatible with the intraspecific variability observed in various
members of Pleurodira (see Georgalis et al. 2020b for
a discussion). Georgalis et al. (2020b) attributed this specimen to
cf. Eocenochelus sp., although it is much likely referable to the same
species as the more complete shell from Medau is Fenus described
above, i.e., Eocenochelus cf. E. eremberti.
Neogene
Locality
San Michele (Municipality of Laerru, Anglona subregion, northern
Sardinia).
Geological setting and age
Perfugas Formation (Early Miocene [early–middle Burdigalian,
between 18.83 ± 0.13 and 18.29 ± 0.13 Ma]; Sowerbutts 2000;
Oudet et al. 2010). The fossiliferous site of San Michele is located
at the northern periphery of the village of Laerru. An epiclastic
pomiceus-cineritic level underlies silicified lacustrine carbonates of
the lower part of the Perfugas Fm. (Sowerbutts 2000). This epiclas­
tic level is divided into three different layers, most of the vertebrate
remains coming from the top of the lower layer (Zoboli and Pillola
2017; Mennecart et al. 2019). In the Anglona subregion, the lower
part of the Perfugas Fm. comprises continental lacustrine deposits
represented by finely bedded limestones and tuffs (Sowerbutts
2000).
Pan-Trionychidae indet.
Figure. 4
Material
MDLCA 14116, undetermined bone fragments and probable costals
(Figure 4).
Description and remarks
The material was originally figured by Zoboli and Pillola (2017)
who treated it as a probable trionychid. MDLCA 14116 is
a rather fragmentary specimen, consisting of few shell and
limb bone fragments on a slab. The largest and most well-

Figure 5. Pan-Trionychidae indet. (MDLCA 23859), Noragugume, Burdigalian (Arenarie di Dualchi Fm.), almost complete nuchal in dorsal (a), ventral (d), anterior (c), and
posterior (d) views.
preserved elements among these is a probable costal fragment.
The element bears a distinctive sculpturing pattern, consisting
of dense connected ridges forming a honeycomb-like pattern.
This fragmentary specimen is little informative, but neverthe­
less, the presence of the distinctive sculpturing pattern along
with the absence of shell scutes denote a referral to Pan-
Trionychidae (Georgalis and Joyce 2017). A more precise allo­
cation within pan-trionychids is not possible.
HISTORICAL BIOLOGY 7
Locality
Nulvi (Anglona subregion, northern Sardinia).
Geological setting and age
The precise location of the fossiliferous locality is unknown and the
only available geological data were published by Portis (1901b) who
reported an internal mould of a carapace collected by Domenico
Lovisato in a clay-limestone block initially referred to the Middle
8 D. ZOBOLI ET AL.
Figure 6. Cheloniidae indet. (MDLCA 23858), Noragugume, Burdigalian (Arenarie di
Dualchi Fm.). Portion of carapace (three partial left costals and two neurals) in
dorsal view (a) and interpretative drawing (b).
Miocene. Littoral carbonates from the second sedimentary cycle of
Sardinia crop out in the area of Nulvi and are referable to the late
Burdigalian (Carmignani et al. 2008).
?Pan-Trionychidae indet.
Material
The internal mould of a carapace probably destroyed during the
WWII when the former geology institute of Cagliari was bombed.
Description and remarks
Portis (1901b) described a carapace from Nulvi which he referred to
Procyclanorbis sardus. However, he did not provide any figure of
this specimen, which is currently lost, probably destroyed during
WWII. Portis (1901b) reported that the specimen from Nulvi was
similar but smaller than the holotype of Procyclanorbis sardus (see
locality Is Mirrionis-Piazza d'Armi).
Locality
Noragugume (Marghine subregion, central Sardinia).
Geological setting and age
Arenarie di Dualchi Formation (Early Miocene [Burdigalian],
Porcu 1983). The remains of two different turtle specimens
originate from a coarse sandstone of fluvial or transitional
environment that just precedes the Burdigalian marine trans­
gression, cropping out along the SP33, about 250 m N of the
village of Noragugume. All fossils were collected associated
together in the same layer in which bone remains of indetermi­
nate artiodactyls were also found.
Pan-Trionychidae indet.
Figure. 5
Material
MDLCA 23859 (formerly CL 151), an almost complete nuchal
(Figure 5).
Description and remarks
MDLCA 23859 represents an almost complete nuchal. It is
a mediolaterally elongated element, 180 mm wide and 56 mm
long with a semi-elliptical shape. Its anterior margin is relatively
straight (symmetrically wavy), while its posterior one is broadly
convex. In that posterior margin, there is a recognisable small
medial concavity, a sinuosity to which the neural I was sutured.
The visceral surface shows in the central area a weak ridge trans­
versally elongated and, posterior to it, a depression. Distinct sculp­
turing covers the dorsal surface of the nuchal, consisting of dense
connected and disconnected ridges. This sculpturing is more pro­
minent in the posterior portion of the bone and slightly fades out
towards its anterior margin, while at its lateral edges it consists
mainly of a honeycomb-like pattern, with thin connected ridges.
The presence of the distinctive sculpturing pattern along with the
absence of shell scutes permit the referral of specimen to pan-
trionychids (Vitek and Joyce 2015; Georgalis and Joyce 2017).
Interestingly, the nuchal is much different than that of Trionyx
Geoffroy Saint-Hilaire, 1809, the most common and abundant
genus of trionychids in the Neogene of Europe (Georgalis and
Joyce 2017); moreover, it appears to be also different than that of
Rafetus Gray, 1864, which is the sole other so far recognised genus
in the Neogene of Europe, though still much rarer than Trionyx
(Georgalis and Joyce, 2017). In contrast, the straight anterior mar­
gin of the Noragugume nuchal bears some general resemblance to
cyclanorbines (see Figure 6 in Meylan et al. 1990) but other car­
apace and, in particular, plastron elements are required to confirm
or reject any cyclanorbine affinities. In any case, the so far total
absence of pan-cyclanorbines from the Neogene of Europe
(Georgalis and Joyce 2017) could hint for pan-trionychine affinities
for the Noragugume specimen, however, based only on this single
nuchal, it is impossible to make a more precise identification. As
such, the nuchal from Noragugume can only be assigned to as Pan-
Trionychidae indet.
Cheloniidae indet.
Figure. 6
Material
MDLCA 23858 (formerly CL 152), a fragment of a carapace from
a single individual (Figure 6).
 HISTORICAL BIOLOGY 9

Figure 7. Trachyaspis lardyi Meyer, 1843 (MSNM V1599), Bosa, late Burdigalian–late Langhian. Internal mould of a large carapace plus some fragments of the actual
carapace showing their external surface (a) and interpretative drawing (b).

Description and remarks

MDLCA 23858 consists of a portion of a large sized shell with
a smooth surface. It comprises three partial left costals and two
neurals. Although the remain is heavily fractured (in its present
condition, it is the result of the union of several fragments), sutures
among the skeletal elements are clearly visible, as well as are the sulci
among the pleural and vertebral scutes. The best preserved costal
shows nearly parallel anterior and posterior edges, with a modestly
concave anterior edge (and convex posterior one) suggesting that
this fragment likely originates from the left anterolateral portion of
the carapace. Both the neurals are clearly hexagonal with shorter
anterolateral edges. The costals can be tentatively identified as the
three first left costals (the first one is only minimally preserved) and
the neurals as the second and third elements of the series. The sulci
delimiting the vertebrals indicate that the general shape of these
scutes was rather rectangular and not distincty hexagonal.
The general morphology of these remains fits well with that of
the carapace of members of Cheloniidae and the general size as well
the nearly rectangular vertebrals indicate that they belonged to
a morphologically mature individual (among others, see Wyneken
2001; Kordikova 2002). The material is not informative enough for
a more precise identification but the broad, nearly rectangular
vertebrals and the position of the interpleural sulci (nearly perpen­
dicular to the carapace sagittal axis) allow us to tentatively exclude
the genera Eretmochelys Fitzinger, 1843, and Lepidochelys, 1843,
among the genera that were or are still present in Europe
(Młynarski 1976; Lapparent de Broin, 2001; Kordikova 2002;
Speybroeck et al. 2020).

Figure 8. Trachyaspis lardyi Meyer, 1843 (MSNM V1599), Bosa, late Burdigalian–late Locality
Langhian. Detail of the scultpuring pattern of the right costals. Bosa (Planargia subregion, central-western Sardinia).
10 D. ZOBOLI ET AL.
Figure 9. ?Testudinoidea indet., Fiume Santo (Porto Torres), Tortonian–early
Messinian. Right xiphiplastron (FS1995#0579) in ventral (a) and dorsal (b) views.
Geological setting and age
The geological data and the precise provenience of the fossil are not
available. The Miocene succession in the Bosa area is represented by
marls, sands, and conglomerates from the second marine sedimen­
tary cycle of Sardinia (Early–Middle Miocene [late Burdigalian–late
Langhian]) (Carmignani et al. 2008).
Trachyaspis lardyi Meyer, 1843
Figures. 7–8
Material
MSNM V1599, an internal mould of a carapace with fragments of
the left and right costal III and right costal IV (Figures 7–8).
Description and remarks
MSNM V1599 represents the internal mould of a large carapace
(preserved length and width: 480 and 450 mm respectively) plus
some fragments of the actual carapace showing its external
surface (Figures 7–8). The carapace gets significantly narrower
towards its posterior margin. The mould of the carapace pre­
serves the imprints of neurals III–VII (VIII?), most of the
costals (all except I), and perhaps also the pygal; less clear are
the possible boundaries of suprapygals I–II, (Figure 7). The
Figure 10. Cheloniidae indet. (MDLCA 23860), Sassari, probably Monte Santo Fm. (Tortonian–early Messinian). Partial mould of the visceral surface of a carapace and
fragments of costals (a) and interpretative drawing (b).
neurals are elongated and hexagonal, with their anterior margin
being much shorter than the posterior one. The surface of the
mould of some costals shows the presence of a furrow from the
centre towards the distal region of the bony elements represent­
ing the position of the internally swollen rib. The few remains
of the actual carapace reveal the presence of a distinctive scult­
puring pattern on the shell, consisting of dense ridges
(Figure 8). Based on this sculpturing pattern, the presence of
shell scutes, and the overall shape of the carapace, we refer the
specimen to the cheloniid Trachyaspis Meyer, 1843, and its only
recognised valid species Trachyaspis lardyi Meyer, 1843 (see
among others Chesi et al. 2007b; Villa and Raineri 2015).
Locality
Fiume Santo (Municipality of Porto Torres, Nurra subregion,
northwestern Sardinia).
Geological setting and age
This is an alluvial deposit (Late Miocene [Tortonian–early Messinian])
cropping out within the industrial complex of local thermoelectric
power station. A few small carapace or plastron fragments were col­
lected in the site of Fiume Santo, in the Upper Miocene alluvial
sediments of the third Miocene sedimentary cycle recognised in
Sardinia. The detailed stratigraphic data and the geological context of
this site are not satisfactorily known; however, a late Tortonian–early
Messinian age is suggested by the associated Turolian mammal fauna
(Abbazzi et al. 2008b; Casanovas-Vilar et al. 2011).
?Testudinoidea indet.
Figure. 9
Material
FS1995#0579, partial right xiphiplastron (Figure 9); FS IV – 139/1,
incomplete pygal; FS1994#7, one shell fragment; FS1994III#258/4, one
shell fragment; FS1994III#257/2, two shell fragments; FS1995III#91,
five small shell fragments; FS30.01.1995RecuperoTrincea, three shell
fragments; FS1995IIITrincea 142–143-144-145-147, five shell
fragments.

Description and remarks
Turtle remains from Fiume Santo are represented by a few shell frag­
ments that have been so far only partly studied. Remarkable is
FS1995#0579 (Figure 9), which has been tentatively considered by
Abbazzi et al. (2008b) to be the postero-lateral portion of a right xiphi­
plastron. There are no preserved sutures and clear scute sulci, but
a transverse ‘step’ on the ventral surface of the bone has been interpreted
as a possible boundary between the femoral and anal shields (Abbazzi
et al. 2008b). The area posterior to the ‘step’ is slightly concave.
The morphology of FS1995#0579 is reminiscent of the xiphi­
plastron of some testudinoids but it is considered too fragmentary
even for a tentative identification beyond Testudinoidea indet.
Should it represent a member of this group it would be the oldest
representative on the island.
Locality
Sassari (north-western Sardinia).
Geological setting and age
Probably Monte Santo Formation (Late Miocene [Tortonian–early
Messinian], Pomesano Cherchi 1971; Mazzei and Oggiano 1990).
The turtle specimen herein described was found in an old piece of
furniture during the relocation work of the former Department of
Geology at the University of Cagliari. The origin area (Sassari) is the
only available data reported on the label associated with the fossil
and the exact site of provenance is unknown. However, based on
the carbonate nature of the sediment, represented by a marine
bioclastic calcarenite, it is possible to conclude that the fossil origi­
nates from the Late Miocene Monte Santo Fm. (Tortonian–early
Messinian) that extensively crops out in the Sassari area.
Cheloniidae indet.
Figure. 10
Material
MDCLA 23860, an incomplete internal mould of a carapace with
a few bone fragments (Figure 10).
Figure 11. Pan-Trionychinae indet. (MDLCA 14007; holotype of Procyclanorbis sardus Portis, 1901b), Is Mirrionis-Piazza d'Armi area (Cagliari), late Tortonian–early Messinian
(Calcari di Cagliari Fm.). Partial carapace (a) and its mould (b). For an interpretative drawing see Georgalis et al. (2017).
HISTORICAL BIOLOGY 11
Description and remarks
The block preserves the mould of the visceral surface of a portion of
a carapace extending from the costal IV to VIII (maximum preserved
width and length are 27.5 and 21.0 cm respectively). Each costal
shows in its central sector a relatively deep, longitudinal groove
corresponding to the rib; these depressions end medially with
a deep hole hosting in some cases a part of the rib head. It is not
possible to detect the imprints of the sutures delimiting the neurals
and the suprapygal/s as well as peripheral elements or the pygal. The
area formerly covered by the neurals shows a deep, sagittal groove
corresponding to the vertebral centra (not preserved). Fragments of
costals are present at the periphery of the mould, both to the left and
to the right. Noteworthy is that the eighth costal rib is pointing
towards the posterior, a feature characteristic of marine turtles
(Parham and Fastovsky 1997; Parham 2005).
The general morphology of MDLCA 23860 is congruent with
that of a cheloniid turtle, particularly in the orientation of the eight
costal ribs, and accordingly we refer it to Cheloniidae.
Locality
Is Mirrionis-Piazza d'Armi, Cagliari (southern Sardinia).
Geological setting and age
Calcari di Cagliari Formation (Late Miocene [late Tortonian–early
Messinian]; Barca et al. 2005). The Calcari di Cagliari Fm. is
represented by three main lithofacies that are, from the bottom to
top: the ‘Pietra Cantone’, the ‘Tramezzario’ and the ‘Pietra Forte’
(Gandolfi and Porcu 1967; Cherchi 1974). The bottom lithofacies
(‘Pietra Cantone’) is represented by yellow marly-arenaceous lime­
stone with common intense bioturbation. The ‘Tramezzario’ is
represented by calcarenites and marls with abundant bioclastic
components. This lithofacies shows widespread phenomena of
synsedimentary breccias, slumpings, erosional surfaces and fault­
ing. The uppermost lithofacies (‘Pietra Forte’) consists of massive
coarse biostromal limestones (Barca et al. 2005). The carbonate
succession of Cagliari is of a Late Miocene age, with the
Tortonian–Messinian boundary tentatively placed within the
upper part of the ‘Pietra Cantone’ (Georgalis et al. 2017). The turtle
remains from Is Mirrionis-Piazza d'Armi originate from the
‘Tramezzario’ lithofacies (Portis 1901b; Comaschi Caria 1959).
12 D. ZOBOLI ET AL.

Figure 12. Pan-Trionychinae indet. (MDLCA 14008), Is Mirrionis-Piazza d'Armi area

(Cagliari), late Tortonian–early Messinian (Calcari di Cagliari Fm.). Partial internal
mould in dorsal view.

Pan-Trionychinae indet.

Figures. 11–12

Material
MDLCA 14007 (holotype of Procyclanorbis sardus Portis, 1901b)
(Figure 11), an incomplete carapace and its internal mould;
MDLCA 14008, the internal mould of a carapace (Figure 12).

Remarks
The first pan-trionychid find from the area, the incomplete shell
MDLCA 14007 (Figure 11) was described as a new species of
a supposed cyclanorbine pan-trionychid, Procyclanorbis sardus, by
Portis (1901b). Subsequent finds of pan-trionychids from the same
locality were later made by Comaschi Caria (1959). Georgalis and
Joyce (2017) and Georgalis et al. (2017) casted doubt on the validity of
Procyclanorbis sardus and its purported affinities with cyclanorbines
and instead treated it as an indeterminate pan-trionychine. For
Figure 13. Cheloniidae indet., Is Mirrionis-Piazza d'Armi area (Cagliari), late
a detailed description of the holotype of Procyclanorbis sardus and Tortonian–early Messinian (Calcari di Cagliari Fm.). Cranium in palatine view (a)
the material of Comaschi Caria (1959), see Georgalis et al. (2017). and a partial left hyo-hypoplastron (b). These specimens are currently lost and were
MDLCA 14008 (Figure 12) is a mould of the visceral surface of originally referred to Procyclanorbis sardus by Portis (1901b).
an incomplete carapace. The fossil was originally described and
assigned to Amyda sardus by Comaschi Caria (1959), and more
recently briefly redescribed and figured by Georgalis et al. (2017). shell. Currently, large parts of the carapace imprint are missing and
Since its original description, the specimen has suffered severe the sutures between most costals have faded. Based on the absence
degradation, although it was initially an almost complete imprint of peripherals and shell scutes, Georgalis et al. (2017) considered
of a carapace, missing only its upper right and lower margins of the this specimen to be an indeterminate pan-trionychine.
 HISTORICAL BIOLOGY 13

Description and remarks

Both specimens were originally referred by Portis (1901b) to the
pan-trionychid Procyclanorbis sardus. Nevertheless, they have since
been suggested to belong to cheloniids instead (Broin de 1977;
Kotsakis 1985; Georgalis and Joyce 2017; Georgalis et al. 2017).
For more details, see Georgalis et al. (2017).

?Testudines indet.

Figure. 14

Material
Repository unknown, undetermined bone element.

Figure 14. ?Testudines indet. The specimen is currently lost and was originally
referred to as an iliac bone of Trionyx by Comaschi Caria (see pl. I, Figures 3-4 in
Comaschi Caria 1959).
Remarks
The fossil has been illustrated and uncertainly identified as an iliac
bone of Trionyx by Comaschi Caria (1959). This attribution was
exclusively based on the similarity with the iliac bones of Trionyx
pliocenicus illustrated by Fucini (see pl. V, Figure 5 in Fucini 1912).
The fossil was not found in the collection of MDLCA, therefore its
redescription is not possible.

Locality
Cheloniidae indet. Sant'Avendrace, Cagliari.

Geological setting and age

Figure. 13 Calcari di Cagliari Formation (Late Miocene [late Tortonian–early
Messinian]). The turtle from Sant'Avendrace originates from the
‘Tramezzario’ lithofacies (see above) and was found in the garden of
Material the nunnery ‘Congregazione delle Ancelle Della Sacra Famiglia’.
A cranium (Figure 13a) and a partial left hyo-hypoplastron The fossil is currently embedded in a fountain inside the garden of
(Figure 13b), probably destroyed during WWII. the nunnery (Zoboli et al. 2021).

Figure 15. Cheloniidae indet. (unnumbered, the specimen is set in a fountain inside the garden of the nunnery ‘Congregazione delle Ancelle della Sacra Famiglia’),
Sant'Avendrace (Cagliari), late Tortonian–early Messinian (Calcari di Cagliari Fm.). Internal mould of a carapace with few bone fragments (a) and interpretative drawing (b).
14 D. ZOBOLI ET AL.
Figure 16. Cheloniidae indet. The now lost carapace fragment of the specimen
from Sant'Avendrace according to Comaschi Caria (1959: pl. I, Figure 2).
Cheloniidae indet.
Figures. 15–16
Material
Unnumbered, the incomplete internal mould of a carapace with
some bone fragments (Figures 15–16).
Description and remarks
Comaschi Caria (1959) described this shell and referred it to
Amyda burdigalensis (Bergounioux, 1935), a pan-trionychid spe­
cies originally known from the early Oligocene of France that is
currently regarded to be a nomen dubium (see Georgalis and
Joyce 2017). Zoboli and Pillola (2016b) regarded this specimen
to represent instead a cheloniid, a view that was followed also
by Georgalis and Joyce (2017). The specimen is apparently
indeed a cheloniid, judging from the posterior orientation of
the eighth costal rib (see Parham and Fastovsky 1997; Parham
2005). A carapace fragment from the same shell figured by
Comaschi Caria (1959) (Figure 16), and now lost, bears
a distinctive ornamentation, which could hint towards affinities
with the cheloniid genus Trachyaspis (but we refrain to identify
it at genus level without having the possibility of directly eval­
uating any diagnostic character).
Locality
Nuraghe Su Casteddu (Municipality of Dorgali, Supramonte sub­
region, central-eastern Sardinia).
Geological setting and age
Nuraghe Casteddu Formation (Pliocene [Piacentian]; Massari
and Dieni 1973). The Nuraghe Casteddu Fm. is an alluvial
succession principally represented by sand and gravel deposits
and subordinate pelitic levels with gastropod and rare vertebrate
remains. The vertebrate taxa reported in the locality of Nuraghe
Su Casteddu are: Discoglossus sp., Testudinoidea indet. (for­
merly Emydidae indet.), cf. Archaeolacerta sp., ‘Colubrinae’
indet., Talpa sp., Chiroptera indet., Asoriculus aff.
A. gibberodon, and Tyrrhenoglis cf. T. majori (Esu and
Kotsakis, 1980; Zammit Maempel and de Bruijn 1982, 1983;
Sanchiz 1998; Delfino et al. 2011).
Testudinoidea indet.
Material
Repository unknown, a very small fragment of a carapace.
Description and remarks
Esu and Kotsakis (1980, 1983) reported a very small fragment
of a carapace, which they assigned to Emydidae indet. The
authors did not provide any description or figure of the fossil
material. We further note that the taxonomic concept and
content of Emydidae has substantially changed since the
1980ʹs and this name is currently used to define a much
more confined group of aquatic testudinoids (see Joyce et al.
2021). Apparently, Esu and Kotsakis (1980, 1983) intended to
refer this material to an aquatic testudinoid – however, on the
absence of any figure or rigorous description, we cannot more
precisely refer it to either emydids or geoemydids, and there­
fore we must consider it as an indeterminate testudinoid.
Quaternary
Locality
Capo Mannu D4 Local Fauna (Municipality of San Vero Milis, Sinis
Peninsula, central-western Sardinia).
Geological setting and age
Capo Mannu Formation, D4 Local Fauna (Early Pleistocene)
(Pecorini et al. 1974). The fossil tortoise originates from a Late
Pliocene–Early Pleistocene coastal dune complex, precisely
from the base of D4 dune, originally referred to the Late
Pliocene (Carboni and Lecca 1995; Abbazzi et al. 2008a) and
currently considered to be Early Pleistocene in age (Kotsakis
et al. 2008b). The associated vertebrate taxa include the dwarf
suid Sus sondaari Van der Made, 1999, the bovid Nesogoral
sp., and Bovidae gen. et sp. indet. (large and small-sized
forms).
Testudo pecorinii Delfino in Abbazzi et al. 2008a
Figure. 17
Material
MDLCA 23789 (formerly CL 102), partial carapace and remains of
plastron (Figure 17).
Description and remarks
This taxon is based on a single and incomplete shell filled with
arenaceous matrix (Abbazzi et al. 2008a). The specimen shows
well-preserved skeletal elements but the bone surface displays only
few anatomical details. The shell has a total length of 22.5 cm,
a maximum width of 17.8 cm, and a maximum height of
12.2 cm. The best-preserved area of the carapace is the medial
part, whereas the anterior and posterior portions are respectively
eroded and incomplete (e.g., the posterior peripherals are only
partly preserved). The carapace is characterised by five evident
bosses sagittally aligned on the carapace and developed in corre­
spondence to the central region of the relative vertebral scutes.
These bosses are medio-laterally elongated, apically convex, and
well separated from each other (therefore dissimilar from the
pathological condition known as ‘pyramiding’; see for details
 HISTORICAL BIOLOGY 15

Figure 17. Testudo pecorinii Delfino in Abbazzi et al. 2008a (MDLCA 23789, formerly CL 102), Capo Mannu D4 local fauna, Capo Mannu (San Vero Milis), Early Pleistocene.
Holotype, almost complete carapace and plastron in dorsal (a), ventral (b), left lateral (c), anterior (d), and posterior (e) views. For an interpretative drawing see Abbazzi
et al. (2008a)

Abbazzi et al. 2008a). The vertebral area is mediolaterally rather hypoplastra as observed in Testudo. graeca Linnaeus, 1758 and in
broad, broader than in extant Testudo. hermanni Gmelin, 1789. other tortoise taxa with a hinge (Abbazzi et al. 2008a). Similarly
The plastron is incomplete. It is anteriorly bent upwards and to certain other mammalian co-inhabitants in Capo Mannu,
shows posteriorly the separation of the xiphiplastra from the T. pecorinii probably represents an insular endemic form.
16 D. ZOBOLI ET AL.
Figure 18. Testudo hermanni Gmelin, 1789 (MT-VII-must-T-01), Cava VII-must of Monte Tuttavista (Orosei), Early Pleistocene. Incomplete shell in dorso-lateral right view (a)
and interpretative drawing (b).
Locality
Cave VII – quarry ‘mustelide’ fissure filling of Monte Tuttavista
(Municipality of Orosei, Baronie subregion, central-eastern
Sardinia).
Geological setting and age
The Upper Jurassic carbonate complex of Monte Tuttavista is
characterised by a developed karst network rich in fissure fill­
ings, which contain an impressive amount of Quaternary verte­
brate remains (Rook et al. 2003; Abbazzi e al., 2004; Delfino
et al. 2008, 2018; Tschopp et al. 2018; Georgalis et al. 2019;
Angelone et al. 2020). Turtle remains have been reported exclu­
sively for Cava VII-must (Testudo cf. T. hermanni) and VII-
giacimento 2 (Chelonii indet.; see identifications in Rook et al.
2003; Abbazzi et al. 2004). Here we focus on the most informa­
tive remains only, those from Cava VII-must. Associate fauna
includes the insular endemic mammal taxa Macaca majori
(Azzaroli, 1946), Rhagapodemus minor (Brandy, 1978),
Prolagus figaro López Martínez in López Martínez and Thaler,
1975, Pannonictis baroniensis Rook et al., 2018, and Nesogoral
spp. The faunal assemblage suggests that this fissure is one of
the oldest of Monte Tuttavista. In particular, the presence of the
bovid Nesogoral Gliozzi and Malatesta, 1980 is what relates the
fissure to the oldest of the two Sardinian Faunal Complexes
(i.e., Nesogoral FC, Capo Figari/Orosei 1 Faunal sub-Complex),
which can be referred to the Early Pleistocene (Palombo 2018).
Testudo hermanni Gmelin, 1789
Figures. 18–19
Material
MT-VII-must-T-01, an incomplete shell partially embedded in
a block (Figures 18–19a) in association with a fragmentary left
humerus (Figure 19b).
Description and remarks
The incomplete tortoise shell MT-VII-must-T-01 is partly embedded
in a block of rock (Figure 18), with just part of the anterior plastral
lobe isolated from the matrix and removable (Figure 19a). The
carapace shows part of the external surface allowing the detection
of parts of neurals III–VII. Neural VI is clearly hexagonal with the
anterolateral sides only slightly shorter than the posterolateral ones
(configuration 6A of Pritchard 1988). On the right side of the car­
apace, portions of costals I–VII are preserved, whereas on the left
side, only fragments of costals IV–VI are visible. The costals are
trapezoidal and those from III to VI clearly show distal (and also
proximal) ends alternatively broad and narrow. Most of the periph­
eral elements of the right side are at least partly preserved (at least
peripherals II–VIII). The scute sulci are not clearly visible except for
some pleuromarginal sulci overlapping the costopleural sutures.
 HISTORICAL BIOLOGY 17

Figure 19. Testudo hermanni Gmelin, 1789 (MT-VII-must-T-01), Cava VII-must of Monte Tuttavista (Orosei), Early Pleistocene. a, anterior plastral lobe in dorsal (visceral) view
(a1), interpretative drawing of the dorsal view (a2), ventral view (a3), interpretative drawing of the ventral view (a4), lateral left view (a5), and distal view (a6). b, proximal
portion of left humerus in anterior (b1), ventral (b2), and proximal (b3) views.

A 40 mm long and 58 mm wide portion of the anterior plastral lobe
is preserved as a single isolated fragment (Figure 19a). The morphology
of this fragment is very well preserved. The two epiplastra, part of the
entoplastron and a small anterior portion of the hyoplastra are pre­
served. The anterior lobe is ventrally flat and when complete it had
probably a vaguely trapezoidal shape in dorsal or ventral view. Its
anterior rim is slightly roundish, with just a modest sagittal notch. In
dorsal or ventral view, on both sides, there is no (anterolateral) notch in
correspondence of the gularohumeral sulcus. The anteroposterior
development of the epiplastral lips is rather uniform because the area
delimited by the dorsal fold of the gulars shows only a modest medial
concavity. Therefore, the lips are on the whole nearly rectangular in
dorsal view. The lips do not significantly overhang the epiplastra and
do not reach (and therefore do not overhang) the entoplastron. There
is no significant gular pocket. The entoplastron is not complete but was
likely triangular and anteroposteriorly elongated in dorsal view, and
rather roundish in ventral view. The gulars are distinctly elongated
(each 24 mm long and 15 mm wide) and deeply enter the entoplastron;
18 D. ZOBOLI ET AL.
Figure 20. cf. Testudo sp. (MDLCA 23861a-c), Middle Pleistocene, Punta del Quadro
(Alghero). a (MDLCA 23861b), incomplete right C6 in dorsal (a1) and visceral (a2) views;
b (MDLCA 23861c), fragment of a hypoplastron in ventral (b1) and visceral (b2) views;
c (MLDCA 23861a), incomplete left humerus in anterior (c1), dorsal (c2), and posterior
(c3) views, and external imprint and part of the bone embedded in the bone breccia (c4).
their ventral surface is flat and not elongated. On the absence of the
posterior area of the entoplastron, it is not possible to evaluate if the
humeropectoral sulcus entered the entoplastron or not.
The left humerus is represented by the proximal portion only
(Figure 19b). It is sigmoid shaped in anterior and posterior
views. The medial process is significantly taller than the lateral
process and than the articulation surface (or head). In proximal
view, the latters are parallel and delimit a narrow intertubercu­
lar fossa.
Despite its incompleteness, MT-VII-must-T-01 can be referred to
Testudinidae and in particular to Testudo hermanni on the basis of
the following characters (Amiranashvili 2000; Hervet 2000; Delfino
et al. 2012; Luján et al. 2016; Vlachos and Rabi 2018): trapezoidal
costal elements with ends alternatively broad and narrow; overlap of
the pleuromarginal sulci with the costoperipheral sutures; ventral
gular surface flat; epiplastral lips not particularly high and not over­
hanging posteriorly (they do not overhang the entoplastron) so that
they do not develop a significant gular pocket.
Locality
Punta del Quadro, Porto Conte Bay (Municipality of Alghero,
north-western Sardinia).
Geological setting and age
Palombo et al. (2008, 2017) described a fossiliferous succession in
a small marine cave that opens slightly above sea level into the
Jurassic limestone of the Punta del Quadro cliff. The authors inter­
preted the succession as a filling of a palaeo-cave and recognised two
distinct bone breccias. The lower breccia was attributed to the pre-
Tyrrhenian (Middle Pleistocene), while the upper breccia to the post-
Tyrrenian (Late Pleistocene). The tortoise fossils herein described
originate from the pre-Tyrrhenian breccia. Associate fauna includes
insular endemic micromammal taxa: the lagomorph Prolagus sardus
(Wagner, 1829), the rodents Rhagamys orthodon (Hensel, 1856) and
Microtus (Tyrrhenicola) henseli (Major, 1905), and the eulipotyphlan
Asoriculus sp. The faunal assemblage suggests that this breccia is
related to the youngest of the two Sardinian Faunal Complexes (i.e.,
Microtus [Tyrrhenicola] FC, Dragonara Faunal sub-Complex), which
dates to the Middle Pleistocene (Palombo 2018).
cf. Testudo sp.
Figure. 20
Material
MDLCA 23861a, an incomplete left humerus (Figure 20c); MDCLA
23861b, a possible incomplete right C6 (Figure 20a); MDCLA
23861c, a possible fragment of a hypoplastron (Figure 20b). All
the remains likely belong to the same individual.
Description and remarks
MDCLA 23861b (Figure 20a) is the distal portion of a trapezoidal
costal (probably a right C6) distinctly tapering medially and char­
acterised by an interpleural sulcus located approximately in the
middle of the bone, marked growth marks on the external surface
in proximity of the lateral edge, and a costal process (apically
broken) well expressed on the visceral surface and laterally protrud­
ing. MDLCA 23861c (Figure 20b) is a possible fragment of hypo­
plastron. The incomplete left humerus (MDLCA 23861a) is broken
in two parts, one of which is represented by the incomplete prox­
imal end and shaft (Figure 20c). The head has elliptical shape, and
the shaft is markedly curved. The other part of the bone is partially
embedded in the sediment and consists in part of the proximal and
distal ends (visible in longitudinal sections), and in a small fragment
of the shaft and its external imprint (Figure 20c).
Locality
Monte San Giovanni (between the town of Iglesias and the village of
Gonnesa, Iglesiente subregion, south-western Sardinia).
Geological setting and age
As reported in the associated label at NMB, the sample originates
from a bone breccia at Monte San Giovanni. This site is mentioned
in several papers since the end of the 19th Century (e.g., Major 1882;
Lydekker 1891b, 1901, 1905; Bate 1945), but the location of the
fossiliferous deposit is currently unknown. The recognised micro­
mammals (e.g., Microtus [Tyrrhenicola] henseli [Major, 1905] and
Talpa tyrrhenica; Bate, 1945) indicate a Middle–Late Pleistocene
age; however, an older age cannot be excluded.

Figure 21. Testudinidae indet. (giant size) (NMB Ty. 11544), Monte San Giovanni (Iglesiente), probably Middle–Late Pleistocene. Almost complete ungual phalanx in
anterior (a), posterior (b) and lateral (c-d) views.
Testudinidae indet. (giant size)
Figure. 21
Material
NMB Ty. 11544, an almost complete ungual phalanx (Figure 21).
Description and remarks
NMB Ty. 11544 from Monte San Giovanni represents a very large
(width 1.5 cm, total length 3.4 cm) an almost complete ungual
phalanx. The surface of the bone is almost totally free of sediment,
except for a very-small carbonate encrustation still present the
ventral-proximal side. This detail seems to confirm that the fossil
was found in a karst fissure filling or in a cave context. The phalanx
is elongated, flattened, and relatively narrow, measuring 3.4 cm in
length. It has a weakly concave proximal surface and host on each
side a moderate lateral keel. As reported in the label, Major referred
the fossil to a ‘Phalanx iii pedis’ of ‘Testudo spec.’ (Zoboli 2017). The
Sardinian phalanx bears indeed much resemblance with that of
large to giant-sized testudinids (e.g., the extant Aldabrachelys
Loveridge and Williams, 1957 [pers. observ.] and the Miocene
Titanochelon schafferi [Szalai, 1931]; see Figures 7, 8 in Vlachos
et al., 2020b). Despite the poor nature of the material, the specimen
from Monte San Giovanni testifies for the presence of a very large
tortoise in the Sardinian insular ecosystem during the Pleistocene.
Locality
San Giovanni di Sinis (Municipality of Cabras, Sinis Peninsula,
central-western Sardinia).
HISTORICAL BIOLOGY 19
Geological setting and age
The remains of freshwater turtles from this locality were collected
from the most representative Late Pleistocene (Calamosca sub­
synthem, Tyrrhenian, MIS 5e) outcrops of San Giovanni di Sinis
(Caloi et al. 1980; Carboni and Lecca 1985; Lecca and Carboni 2007;
Chesi et al. 2007a). In particular, the fossils originate from the beds
M2d of the ‘Phoenician graves’ section, represented by fine silty sand,
carbonatic silt, carbonatic mud, and marly-sandy limestone. The beds
testify even episodic emersion phases (Carboni and Lecca 1985; Lecca
and Carboni 2007). The associated fauna includes a ‘colubrine’ snake,
the rodent Microtus (Tyrrhenicola) henseli (Major, 1905), the lago­
morph Prolagus sp. (the age suggests P. sardus [Wagner, 1829]), and
the megacerine cervid Praemegaceros cazioti (Depéret, 1897) (Caloi
et al. 1980; Chesi et al. 2007a).
?Emys orbicularis (Linnaeus, 1758)
Material
Collection unknown, three epiplastra, two xiphiplastra, and many
isolated peripherals.
Description and remarks
Caloi et al. (1980) briefly described turtle remains from San
Giovanni di Sinis, which they referred to the extant Emys orbicularis
(Linnaeus, 1758). On the absence of any published figure, however,
it is not possible to confirm this identification, especially when
considering that the same authors figured from the same locality
another aquatic testudinoid, the geoemydid Mauremys (Caloi et al.,
1980). Therefore, the presence E. orbicularis at San Giovanni di
Sinis has to be considered doubtful, even if its identification on the
basis of easily identifiable, diagnostic elements (epiplastra and
xiphiplastral) should be correct.
20 D. ZOBOLI ET AL.

Figure 22. Mauremys sp., San Giovanni di Sinis (Cabras), Late Pleistocene (Calamosca subsynthem, Tyrrhenian, MIS 5e). a (MDLCA 23853, formerly UC 1), fragment of
anterior plastral lobe (epiplastra and entoplastron) in ventral (a1), visceral (a2), and anterior (a3) views; b (MDLCA 23854, formerly UC 2), pygal in dorsal (b1) and ventral
(b2) views; c (MDLCA 23855, formerly UC 3), fragmentary right hyoplastron of a juvenile specimen in ventral (c1) and visceral (c2) views; d (MDLCA 23856, formerly UC 4),
fragmentary left xiphiplastron in ventral (d1) and visceral (d2) views; e (MDLCA 23857a, formerly UC 5), fragment of xiphiplastron in ventral (e1) and visceral (e2) views;
f (MDLCA 23857b, formerly UC 5), fragment of xiphiplastron in ventral (f1) and visceral (f2) views; g (MDLCA 23857c, formerly UC 5), fragment of xiphiplastron in ventral (g1)
and visceral (g2) views; h (MDLCA 23857d, formerly UC 5), fragment of xiphiplastron in ventral (h1) and visceral (h2) views; i (MDLCA 23857e, formerly UC 5), fragment of
xiphiplastron in ventral (i1) and visceral (i2) views.

Mauremys sp. a femoro-anal sulcus running perpendicularly to the lateral edge

Material
MDLCA 23851, three plastron fragments; MDLCA 23852, twenty­
three shell fragments; MDLCA 23853 (formerly UC 1),
a fragmentary of an anterior plastral lobe (epiplastra and entoplas­
tron) (Figure 22a); MDLCA 23854 (formerly UC 2), a pygal
(Figure 22b); MDLCA 23855 (formerly UC 3), fragmentary right
hyoplastron of a young specimen (Figure 22c); MDLCA 23856
(formerly UC 4), a fragmentary left xiphiplastron (Figure 22d);
MDLCA 23857a–e (formerly UC 5), five fragments of xiphiplastra
(Figure 22e–i). In addition, Caloi at al. (1980) reported the follow­
ing material kept in Rome: a left peripheral I; a right hyoplastron;
a left xiphiplastron; a fragment of hypoplastron.
Description and remarks
Geoemydid material from San Giovanni di Sinis was initially
described by Caloi et al. (1980) who referred it to Mauremys cf.
caspica (Gmelin, 1774). Chesi et al. (2007a, 2008) subsequently
described more geoemydid remains from the same locality and
referred it to an indeterminate species of Mauremys, because extant
species cannot be identified based on their shell morphology.
Among others, typical of Mauremys are the xiphiplastra with
of the bone and delimiting a broad and deep anal notch; pygal much
broader than long with a longitudinal sulcus (corresponding to the
contact between the two last marginal shields) but without any
transversal sulcus (corresponding to the contact among the fifth
vertebral and the two last marginal shields).
Locality
Grotta Corbeddu (685 SA/NU, Municipality of Oliena, Supramonte
subregion, central-eastern Sardinia).
Geological setting and age
Sondaar et al. (1984) reported the remains of Emys orbicularis in the
layer 3 of the main pit in hall 2 of the cave (latest Pleistocene). This
layer consists of a homogeneous red clay extremely rich in fossils of
the deer Praemegaceros cazioti.
?Emys orbicularis (Linnaeus, 1758)
Material
Unknown.
 HISTORICAL BIOLOGY 21

Figure 23. Testudinoidea indet., Grotte di Is Zuddas (Santadi), Middle–Late Pleistocene. a (MDLCA 23862), costal fragment in ventral (a1), visceral (a2), and lateral (a3)
views; b (MDLCA 23863), nearly complete right humerus in dorsal (b1), anterior (b2), ventral (b3), posterior (b4), and proximal (b5) views; c (MDLCA 23865), incomplete
right femur in posterior (c1), medial (c2), anterior (c3), and lateral (c4) views; d (MDLCA 23864), right femur in posterior (d1), medial (d2), anterior (d3), lateral (d4), and
proximal (d5) views.

Remarks
The material from this locality has not been described and figured,
therefore it is impossible to verify if the fossils are referable to Emys
orbicularis. This taxon has not been retrieved in the 2009 field
season lead by the Soprintendenza per i Beni Archeologici per le
province di Sassari e Nuoro (M. Delfino pers. obs.).
Locality
Grotte di Is Zuddas (763 SA/CI, Municipality of Santadi, Sulcis
subregion, south-western Sardinia).
very rich in mammal and bird remains. The mammal association is
clearly referable to the Microtus (Tyrrhenicola) Faunal Complex
(Dragonara Faunal sub-Complex) and therefore indicates
a Middle–Late Pleistocene age (Palombo 2018).
Testudinoidea indet.
Figure. 23

Geological setting and age Material

The Grotte di Is Zuddas is the largest tourist cave in the SW MDLCA 23862, costal fragment (Figure 23a); MDLCA 23863,
Sardinia, the karst complex is developed in the lower Cambrian nearly complete right humerus (Figure 23b); MDLCA 23864, right
metacarbonates of Monte Meana (Santadi). Different clastic depos­ femur (Figure 23d); MDLCA 23865, incomplete right femur
its are present in the cave. The turtle fossils come from a deposit (Figure 23c).
22 D. ZOBOLI ET AL.
Figure 24. Emys orbicularis (Linnaeus, 1758) (MARTEL 0068, formerly MPSMC 19), Grotta di Monte Meana (Santadi), Late Pleistocene–?early Holocene. Complete carapace in
dorsal view (a) and interpretative drawing (b).
Description and remarks
MDLCA 23862 is a thin distal fragment of a costal elements
characterised by a uniform thickness, a moderate curvature, and
a longitudinal interpleural sulcus on its external surface. The
right humerus MDLCA 23863 (Figure 23b) is nearly complete
(just the distal end is partly eroded) and 26.0 mm long. The shaft
is moderately curved. The intertubercular fossa is very broad and
separates the lateral and medial processes. The latter is taller than
the humeral head and is proximomedially expanded. The ectepi­
condylar foramen is only partly preserved but its groove clearly
indicates its position. The right femur MDLCA 23864
(Figure 23d) shows an elongated and moderately curved shaft.
The femoral head is much taller than the trochanters. The inter­
trochanteric fossa is ventrally open.
The morphology of the zeugopodia from Grotte di Is Zuddas
easily allows us to exclude their referral to Testudo (among others,
the humeral intertubercular fossa is not broad and the femoral
intertrochanteric fossa is ventrally close), but we are not able to
discriminate between emydids and geoemydids whose limb mor­
phology is rather uniform. The thinness and general morphology of
the costal fragment is congruent with such identification.
Locality
Grotta di Monte Meana (2478 SA/CI, Municipality of Santadi,
Sulcis subregion, south-western Sardinia).
Geological setting and age
The turtle remain was found partially embedded in a carbonate
concretion into the cave ‘Grotta di Monte Meana’, near the village
of Santadi. The absence of associated fauna makes a uncertain
chronological attribution of the deposit however the nature of the
concretion and the cave context suggest a probable Late Pleistocene
age (if not more recent).
Emys orbicularis (Linnaeus, 1758)
Figure. 24
Material
MARTEL 0068, a complete carapace.
Description and remarks
The complete carapace MARTEL 0068 has been described by Chesi
et al. (2008) in a conference proceeding. The carapace is 145 mm long
and 105 mm wide (maximum width at the level of the seventh
peripheral bone) and is practically undistorted. The nuchal plate is
hexagonal, wider than long, with anterolateral sides slightly longer than
the postero-lateral ones. Eight neural plates are present and the neural
series is slightly irregular. The peripheral plates are eleven in number;
they are longer than wide. Eight pleural pairs are present: the first one is
 HISTORICAL BIOLOGY 23

Table 1. Palaeontological and archaeological record of Sardinian turtles.

TAXON LOCALITY (Municipality) AGE (Formation or geological context) REFERENCES
Eocenochelus cf. E. eremberti Medau is Fenus (Carbonia) middle–late? Eocene (Cixerri Fm.) Georgalis et al. (2020b)
cf. Eocenochelus sp. Murecci (Gonnesa) middle–late? Eocene (Cixerri Fm.) Georgalis et al. (2020b)
Trachyaspis lardyi Bosa late Burdigalian–late Langhian Chesi (2008), this work
Caretta caretta Turris Libisonis (Porto Torres) Holocene, Roman Epoch (archaeological context) Wilkens (2003)
Caretta caretta Santa Filitica (Sorso) Holocene, Middle Ages (archaeological context) Wilkens (2003)
Caretta sp. Santissima Trinità di Saccargia Holocene, Middle Ages (archaeological context) Baldino (2000)
(Codrongianos)
Cheloniidae indet. Noragugume Burdigalian (Arenarie di Dualchi Fm.) Chesi (2008), this work
Cheloniidae indet. Sant'Avendrace (Cagliari) late Tortonian–early Messinian (Calcari di Cagliari Comaschi Caria (1959), this work
Fm.)
Cheloniidae indet. (= ‘P. sardus’) Is Mirrionis-Piazza d' Armi (Cagliari) late Tortonian–early Messinian (Calcari di Cagliari Portis (1901b), Georgalis et al. (2017)
Fm.)
Cheloniidae indet. Sassari Tortonian–early Messinian (?Monte Santo Fm.) this work
Emys orbicularis Grotta di Monte Meana (Santadi) Late Pleistocene–?early Holocene (bone breccia) Chesi et a. (2008)
Emys orbicularis Grotta Su Guanu (Oliena) Holocene (Neolithic) Kotsakis (1983)
Emys orbicularis S'Imbalconadu (Olbia) Holocene, Roman Epoch (archaeological context) Delussu (2000)
?Emys orbicularis San Giovanni di Sinis (Cabras) Late Pleistocene, Tyrrhenian MIS 5e (Calamosca Caloi et al. (1980)
subsynthem)
?Emys orbicularis Grotta Corbeddu (Oliena) Late Pleistocene (cave deposit) Sondaar et al. (1984)
Emys sp. Santissima Trinità di Saccargia Holocene, Middle Ages (archaeological context) Baldino (2000)
(Codrongianos)
Mauremys sp. San Giovanni di Sinis (Cabras) Late Pleistocene, Tyrrhenian MIS 5e (Calamosca Chesi et al. (2007a)
subsynthem)
Testudo pecorinii Capo Mannu D4 LF (San Vero Milis) Early Pleistocene (Capo Mannu Fm.) Abbazzi et al. (2008a)
Testudo hermanni VII-must, Monte Tuttavista (Orosei) Early Pleistocene (bone breccia) Abbazzi et al. (2004), this work
Testudo sp. Grotta Filiestru (Mara) Holocene (Middle Neolithic) Delussu (2000)
Testudo sp. Santissima Trinità di Saccargia Holocene, Middle Ages (archaeological context) Baldino (2000)
(Codrongianos)
Testudo sp. Santa Filitica (Sorso) Holocene, Middle Ages (archaeological context) Wilkens (2003)
cf. Testudo sp. Punta del Quadro (Alghero) Middle Pleistocene (bone breccia) this work
Testudinidae indet. (giant size) Monte San Giovanni (Iglesias/Gonnesa) ?Middle–Late Pleistocene (bone breccia) Zoboli (2017), this work
Pan-Trionychidae indet. Monte Sinni Mine (Gonnesa) late Ypresian–early Lutetian (Lignitifero Fm.) this work
Pan-Trionychidae indet. Bacu Abis (Carbonia) late Ypresian–early Lutetian (Lignitifero Fm.) Kotsakis (1985)
Pan-Trionychidae indet. San Michele (Laerru) early-middle Burdigalian (Perfugas Fm.) Zoboli and Pillola (2017), this work
Pan-Trionychidae indet. Noragugume Burdigalian (Arenarie di Dualchi Fm.) Chesi (2008), this work
?Pan-Trionychidae indet. (= Nulvi ?late Burdigalian Portis (1901b), Georgalis et al. (2017)
‘P. sardus’)
Pan-Trionychinae indet. Is Mirrionis-Piazza d' Armi (Cagliari) late Tortonian–early Messinian (Calcari di Cagliari Comaschi Caria (1959), Georgalis et al.
Fm.) (2017)
Pan-Trionychinae indet. (= Is Mirrionis-Piazza d' Armi (Cagliari) late Tortonian–early Messinian (Calcari di Cagliari Portis (1901b), Georgalis et al. (2017)
‘P. sardus’) Fm.)
Testudinoidea indet. Nuraghe Su Casteddu (Dorgali) Piacentian (Nuraghe Casteddu Fm.) Esu and Kotsakis (1980, 1983)
Testudinoidea indet. Grotte di Is Zuddas (Santadi) Middle–Late Pleistocene (cave deposit) this work
?Testudinoidea indet. Fiume Santo (Porto Torres) Tortonian–early Messinian (alluvial deposit) Abbazzi et al. (2008b), this work
Testudines indet. VII-giacimento 2, Monte Tuttavista Late Pleistocene (bone breccia) Abbazzi et al. (2004)
(Orosei)
Testudines indet. Monte Sant'Antonio (Siligo) Holocene, Iron Age (archaeological context) Delussu (2000)
Testudines indet. Santissima Trinità di Saccargia Holocene, Middle Ages (archaeological context) Delussu (2000)
(Codrongianos)
?Testudines indet. Is Mirrionis-Piazza d' Armi (Cagliari) late Tortonian–early Messinian (Calcari di Cagliari Comaschi Caria (1959), this work
Fm.)

the longest, whereas the widest is the fourth; the pleural series becomes
narrower posteriorly to it. Two suprapygal plates are present, even if
the suture between the last neural and the first suprapygal is not visible.
The second suprapygal plate is very large and meets posteriorly the
pygal plate, which is about rectangular, wider than long and shows
a small anal notch. The cervical scute is longer than wide and covers
only ¼ of the nuchal length approximately. The first vertebral scute is
lyre-shaped and is the narrowest among the vertebral series; it partially
covers the nuchal, the first peripheral and pleural pairs, and the first
neural. The second to fifth vertebrals are hexagonal and wider than the
corresponding costals. The scute sulcus between the second and the
third vertebrals is located on the third neural, whereas the fifth neural
hosts the third-fourth vertebral sulcus and the eighth neural the fourth-
fifth one. The fifth vertebral completely covers the first and second
suprapygals and partially the pygal plate. On the latter, the scute sulcus
between the last marginals is also visible.
The carapacial morphology clearly fits the one of Emys orbicu­
laris because of the combination of the following characters: the
hexagonal nuchal plate; the little coverage of the narrow cervical on
the nuchal; the narrow lyre-shaped first vertebral; the first costal
pairs partially covering the nuchal; the costo-marginal scute sulci
on the peripherals and not in correspondence with the pleuro-
peripheral sutures; the location of the posterior sulcus of the fifth
vertebral scute on the pygal plate (and not on the second
suprapygal).
Archaeological record
Turtle remains have been reported in different archaeological con­
texts of the island, from the Neolithic to the medieval period. Emys
orbicularis is present in the Neolithic deposit of Grotta Su Guanu
(103 SA/NU), near Oliena (Kotsakis 1983), while Testudo sp. was
24 D. ZOBOLI ET AL.
reported in the Middle Neolithic deposit of Grotta Filiestru (179
SA/SS, Mara, NW Sardinia) (Levine 1983; Delussu 2000; Wilkens
2003). Testudines indet. was reported in the Iron Age Nuragic
sanctuary of Monte Sant' Antonio (Siligo, NW Sardinia) (Delussu
2000). From Roman times, Emys orbicularis was reported from the
site of S'Imbalconadu (Olbia, NE Sardinia), while Caretta caretta
(Linnaeus, 1758) was reported from Turris Libisonis (Porto Torres,
NW Sardinia) (Delussu 2000; Wilkens 2003). Testudo sp., Emys sp.,
Caretta sp., and Testudines indet. were reported from the late
Middle Age convent of Santissima Trinità di Saccargia
(Codrongianos, NW Sardinia) (Baldino 2000; Delussu 2000).
Finally, Testudo sp. and Caretta caretta were reported from the
medieval site of Santa Filitica (Sorso, NW Sardinia) (Wilkens 2003).
Discussion and conclusions
The Sardinian fossil record of turtles documents the occurrence of
an array of different chelonian lineages, including extinct forms, as
well as representatives of extant genera and species (Table 1). No
Mesozoic fossil turtles are yet known from the island; the earliest
Sardinian chelonians are known from the early Eocene, with their
record being almost continuous up to the Holocene, however, with
a total paucity of remains during the whole Oligocene.
Pan-trionychids, commonly known as soft-shelled turtles, repre­
sent the oldest recorded turtle lineage in Sardinia. They are already
present in the island since the early Eocene, judging from the
material from Monte Sinni mine. Slightly younger material from
the middle Eocene of Bacu Abis documents that pan-trionychids
inhabited the island across at least part of the Eocene. Pan-
trionychids are also present and even more abundant in the
Neogene of Sardinia, known from certain Early and Late Miocene
localities. The absence of Oligocene trionychid remains from
Sardinia does not permit any conclusion on whether these
Miocene forms are direct descendants of the Eocene ones or if the
latter became ultimately extinct and the Neogene occurrences
represent the product of some younger dispersal event(s). This is
further complicated by the fragmentary status of most available
fossil remains, which does not afford any more definite conclusion
about their more precise taxonomic affinities. Indeed, all Paleogene
Sardinian pan-trionychids are poorly known and based on rather
fragmentary material that they cannot be more precisely identified
beyond Pan-Trionychidae indet. The same applies to most of the
Miocene occurrences, except for the holotype of Procyclanorbis
sardus which can be identified as a pan-trionychine (Georgalis
et al. 2017). The holotype of Procyclanorbis sardus (MDLCA
14007) and the specimen MDLCA 14008 represent the youngest
occurrence of pan-trionychids in Sardinia (Late Miocene [late
Tortonian-early Messinian]). This time coincides with the relatively
frequent presence of pan-trionychids in other parts of the
Mediterranean during the Tortonian (Georgalis et al. 2016;
Georgalis and Joyce 2017, 2020a). The group seems to have gone
ultimately extinct from Sardinia during the Late Miocene.
European trionychids eventually became extinct during the
Pliocene – recent sporadic occurrences of Trionyx triunguis
(Forskål, 1775) in the Dodecanese Islands (Greece) simply repre­
sent individuals from the nearby Anatolian coast that venture into
the sea (Taskavak et al. 1999; Corsini-Foka and Masseti 2008).
The next oldest turtle clade from Sardinia is Pleurodira. The
occurrences of this group are represented solely by two shells,
probably referable to the middle–late? Eocene (Georgalis et al.
2020b). The most complete shell among these, originating from
Medau is Fenus, has been referred to as Eocenochelus cf.
E. eremberti, whereas the second, less complete specimen, from
Murecci, was assigned to cf. Eocenochelus sp., although it remains
possible that they both belong to the same taxon (Georgalis et al.
2020b). Eocenochelus was a widespread podocnemidid genus across
Western Europe (Pérez-García et al. 2017a; 2019), and its identifi­
cation in Sardinia has been used to support palaeogeographic
reconstructions, according to which, during the early Paleogene,
the Sardinia-Corsica Massif was rather close to continental Western
Europe (Georgalis et al. 2020b). No post-Eocene Sardinian pleur­
odires are known, a fact in concordance with the general demise of
this group in Europe following the latest Eocene; the youngest
occurrences of pleurodires from the continent are recorded from
the Early Miocene of Greece and Malta (Ristori 1895b; Georgalis
and Kear 2013; Georgalis et al. 2013).
Cheloniids from Sardinia are first recorded in the Early Miocene
of Noragugume. The group is also present in Late Miocene local­
ities. Most cheloniid specimens are represented by incomplete shell
remains, however, a single skull is also known. This sole cranial
material originates from the Late Miocene of Is Mirrionis-Piazza d
'Armi and was originally referred to the pan-trionychid
Procyclanorbis sardus by Portis (1901b); the specimen nevertheless
belongs to a chelonioid instead but is now lost and all that remains
is the original photograph of Portis (1901b), which unfortunately
cannot afford much precise information (Georgalis et al. 2017).
Besides the indeterminate cheloniids from Sardinia, the remains
from the Early–Middle Miocene of Bosa and potentially also the
Late Miocene of Sant'Avendrace can be more precisely identified as
Trachyaspis lardyi. The sculptured cheloniid Trachyaspis is other­
wise known also from several Neogene localities in Europe (see
Chesi et al. 2008; Villa and Raineri 2015), probably including also
other Mediterranean islands, e.g., Malta (holotype of Trionyx meli­
tensis Lydekker, 1891a; see Georgalis and Joyce 2017).
Testudinoids are probably first recorded since the Late Miocene,
known by a potential indeterminate testudinoid from Fiume Santo.
Indeterminate testudinoids are also known from the Early Pliocene
of Nuraghe Su Casteddu, originally referred to emydids by Esu and
Kotsakis (1983). However, the exact affinities of this occurrence
cannot be assessed, as the material has never been figured. Three
extant testudinoid lineages have been identified in the Sardinian
fossil record, i.e., emydids, geoemydids, and testudinids.
Emydids have been recovered from the Late Pleistocene of San
Giovanni di Sinis, represented by the extant Emys orbicularis;
however, this was only briefly described and never figured (Caloi
et al. 1980). In any case, E. orbicularis is otherwise known with
certainty from the Late Pleistocene of the island, as a rather com­
plete shell is here described from Grotta di Monte Meana. The
occurrence of E. orbicularis in the Sardinian Quaternary fossil
record, is particularly interesting, taking into consideration that
the extant populations of this species in the island have been
suggested to be the product of some human-aided prehistoric
introduction (Pedall et al. 2011; Rhodin et al. 2021). Indeed, based
on mt-DNA data, Pedall et al. (2011) demonstrated that there is not
adequate differentiation among the Sardinian and other European
populations of E. orbicularis, particularly from Sicily, Corsica, and
the Italian mainland. The same authors accordingly envisaged that
there occurred probably some extirpation event during the
Pleistocene and then the extant populations of E. orbicularis were
subsequently re-introduced by humans (Pedall et al. 2011). As such,
the supposedly endemic Sardinian subspecies Emys orbicularis
capolongoi Fritz, 1995, to which the extant populations from the
island were assigned (Fritz, 1995) is currently not considered valid
(Fritz et al. 2005a; Pedall et al. 2011; Rhodin et al. 2021). The
completeness of the shell from Grotta di Monte Meana described

herein may afford more potential for elucidating important mor­
phological features in the Sardinian Pleistocene Emys, especially
when considering that Sicily is currently inhabited by another
insular endemic species, i.e., Emys trinacris Fritz, Fattizzo,
Guicking, Tripepi, Pennisi, Lenk, Joger, and Wink 2005a (note,
however, that Speybroeck et al. 2020 have treated E. trinacris as
merely a subspecies of E. orbicularis) that is not identifiable on the
basis of its carapace morphology. In any case, it cannot be certain
whether the Late Pleistocene emydids from Sardinia pertained to
Emys orbicularis or represented an insular, now extinct, taxon.
Geoemydids are known exclusively from the Late Pleistocene
of San Giovanni di Sinis, represented by an indeterminate
species of Mauremys (Caloi et al., 1980; Chesi et al. 2007a).
This material is the sole occurrence of geoemydids in Sardinia,
with the group being absent from the extant herpetofauna of
the island. Moreover, as it was suggested by Chesi et al. (2007a),
the San Giovanni di Sinis Mauremys represents the youngest
occurrence of the genus in north central Mediterranean, witnes­
sing a survivorship potentially favoured by insular isolation.
Indeed, Mauremys is abundant in the Late Miocene of Italy
(Ristori 1895a; Chesi et al. 2009; Colombero et al. 2017;
Georgalis et al. 2020a; Georgalis and Delfino 2021; Villa et al.
2021), becoming gradually rarer during the Pliocene of the area
(Sacco 1889; Portis 1893; Collareta et al. 2020), and then extre­
mely rare during the Early and Middle Pleistocene (Portis, 1893;
Kotsakis 1981; Delfino and Bailon 2000), and ultimately absent
in its extant herpetofauna.
Sardinia currently hosts three extant testudinid species, i.e.,
Testudo graeca Linnaeus, 1758, Bringsøe et al. (2001), and Testudo
hermanni Gmelin, 1789, however, all of them are currently
regarded to be the products of human-aided introduction during
prehistoric times or Antiquity (Sindaco et al. 2006; Rhodin et al.
2021). More particularly, for T. graeca, Vamberger et al. (2011)
demonstrated based on molecular data that the extant populations
of this species in Sardinia are the product of a human-aided intro­
duction from northern Africa during prehistoric or even historic
times. Testudo marginata is the largest extant European tortoise
and has a rather disjunct geographic distribution, encompassing
Greece and nearby southern Albania, plus northern Sardinia (Perez
et al. 2012; Rhodin et al. 2021). The Sardinian population was even
treated by Mayer (1992) as its own subspecies, Testudo marginata
sarda Mayer, 1992, however, that opinion has never met acceptance
(Fritz et al. 2005b; Rhodin et al. 2021). In any case, the Sardinian
T. marginata has been generally long considered to represent the
product of human-aided transportation from the Greek mainland
during Antiquity (Angelini 1899; Mayer 1992; Bringsøe et al. 2001;
Fritz et al. 2005b; Rhodin et al. 2021) and this was recently com­
prehensively demonstrated by Perez et al. (2012) with the aid of
molecular data. Finally, also, based on mt-DNA data, Perez et al.
(2014) suggested that the populations of T. hermanni from Sardinia,
Corsica, and Spain, are all the product of human transportation
from Sicily – such opinion for the Sardinian T. hermanni popula­
tions was further accepted by Zenboudji et al. (2016), who also
suggested a potential earlier extirpation of the species in Sardinia
during the Quaternary. We here describe for the first time fossil
material from Sardinia that can be definitely attributable to
T. hermanni, i.e., from the Early Pleistocene of Monte Tuttavista.
This fully concurs with the fact that T. hermanni was autochtho­
nous, at least during the early Quaternary, and then became ulti­
mately extinct from the island before its human-aided
recolonization by allochthonous populations of the species.
Besides the three extant testudinids, there is evidence for
a high diversity of Sardinian tortoises during the latest
Neogene to early Quaternary, comprising two extinct forms,
HISTORICAL BIOLOGY 25
while also another indeterminate species of Testudo is known
from the Middle Pleistocene of Punta del Quadro. The oldest
among these is the endemic insular species Testudo pecorinii,
known from the Late Pliocene–Early Pleistocene of the Capo
Mannu D4 Local Fauna (Abbazzi et al. 2008a). This bizarre
tortoise, characterised by a probable hypo-xiphiplastral hinge
and prominent bosses on the carapace, is more related to
Testudo graeca and T. marginata than to T. hermanni, but is
still rather distinct from all these extant species. The second
extinct tortoise from Sardinia is documented by a large phalanx
from the Middle–Late Pleistocene of Monte San Giovanni. This
specimen testifies that a large-sized testudinid occurred also in
Sardinia. Neogene and/or Quaternary large to giant-sized tor­
toise have been found across many parts of southern Europe
and northern Africa, also including the Mediterranean Islands
(Leith Adams 1877; Depéret and Donnezan 1890–1897;
Lapparent de Broin, 2002; Georgalis and Kear 2013; Luján
et al. 2014; Pérez-García and Vlachos 2014, 2017; Vlachos
et al. 2020a, 2020b; Georgalis and Delfino 2021; Georgalis
et al. 2021; Valenti et al. 2022). Most of these European main­
land giant forms are members of Titanochelon Pérez-García and
Vlachos, 2014 (e.g., Pérez-García and Vlachos 2014; Pérez-
García et al. 2017b; Vlachos et al. 2020a), whereas those from
northern Africa could potentially belong to the extant
Centrochelys Gray, 1872 (see Georgalis et al. 2021). The affinities
of the Mediterranean Island forms have been considered more
problematic (e.g., Luján et al. 2017; Georgalis and Delfino
2021), but a new genus has been recently proposed for some
of the materials coming from Malta, Menorca, and Sicily
(Valenti et al. 2022). Whether the Sardinian large form pertains
as well to the same genus remains to be addressed only on the
basis of new more complete finds from the island.
Acknowledgments
C. Dal Sasso (Museo Civico di Storia Naturale di Milano) and R. Sardella (Museo
Universitario di Scienze della Terra, Sapienza Università di Roma) are thanked for
information about the collections under their care. DZ was supported by grants P.
O.R. Sardegna F.S.E. 2014-2020 - Asse III “Istruzione e Formazione, Obiettivo
Tematico: 10, Obiettivo Specifico: 10.5” Azione dell’accordo di Partenariato:
10.5.12 “Avviso di chiamata per il finanziamento di Progetti di ricerca - Anno
2017” GLP was supported by the Università di Cagliari CAR Project,
“Paleobiodiversità: strumento di base in biostratigrafia, in paleoecologia e nella
valorizzazione dei beni culturali Geo-Paleontologici” GLG acknowledges funding
from the Ulam Program of the Polish National Agency for Academic Exchange
(PPN/ULM/2020/1/00022/U/00001) and Forschungskredit of the University of
Zurich, Grant no. [FK-20-110]. Field work background for this study at Fiume
Santo and Monte Tuttavista has been carried out under agreements between the
“Soprintendenza Archeologia, Belle Arti e Paesaggio per le province di Sassari
e Nuoro” and the Dipartimento di Scienze della Terra dell’Università degli Studi di
Firenze, and have been supported by the University of Florence (Fondi di Ateneo
to LR), the National Geographic Society (grant #7484-03 to LR), the RHOI
program at University of Berkeley (project NSF-BCS-0321893 to LR). Samplings
at Grotte di Is Zuddas have been carried out under agreements between the
“Soprintendenza Archeologia, belle arti e paesaggio per la città metropolitana di
Cagliari e le province di Oristano e Sud Sardegna” and the Dipartimento di Scienze
Chimiche e Geologiche dell’Università degli Studi di Cagliari. We would like to
thank reviewers A. Čerňanský, W.G. Joyce and À.H. Luján for comments that
greatly improved the quality of the manuscript.
Disclosure statement
No potential conflict of interest was reported by the author(s).
Funding
This work was supported by the National Geographic Society [#7484-03].
26 D. ZOBOLI ET AL.
ORCID
Daniel Zoboli http://orcid.org/0000-0001-6820-9538
Georgios L. Georgalis http://orcid.org/0000-0001-7759-6146
Lorenzo Rook http://orcid.org/0000-0001-8923-5428
Massimo Delfino http://orcid.org/0000-0001-7836-7265
References
Abbazzi A, Angelone C, Arca M, Barisone G, Bedetti C, Delfino M, Kotsakis T,
Marcolini F, Palombo MR, Pavia M, et al. 2004. Plio-Pleistocene fossil
vertebrates of Monte Tuttavista (Orosei, Eastern Sardinia, Italy), an
overview. Rivista Italiana Paleontologia Stratigrafia. 110:681–706. doi:10.
13130/2039-4942/5832.
Abbazzi L, Carboni S, Delfino M, Gallai G, Lecca L, Rook L. 2008a. Fossil
vertebrates (Mammalia and Reptilia) from Capo Mannu (Late Pliocene,
Western Sardinia, Italy) with description of a new Testudo (Chelonii,
Testudinidae) species. Rivista Italiana di Paleontologia e Stratigrafia.
114:119–132. doi:10.13130/2039-4942/6371.
Abbazzi A, Delfino M, Gallai G, Trebini L, Rook L. 2008b. New data on the
vertebrate assemblage of Fiume Santo (North-West Sardinia Italy) and over­
view on the Late Miocene Tusco-Sardinian palaeobioprovince.
Palaeontology. 52:251–425. doi:10.1111/j.1475-4983.2008.00758.x.
Advokaat EL, van Hinsbergen DJJ, Maffione M, Langereis CG, Vissers RLM,
Cherchi A, Schroeder R, Madani H, Columbu S. 2014. Eocene rotation of
Sardinia, and the paleogeography of the western Mediterranean region. Earth
Planet Sci Lett. 401:183–195. doi:10.1016/j.epsl.2014.06.012.
Agus M, and Pecorini G. 1978. Livelli a Carofite nel carbone della “prima vena”
della miniera di Seruci e nel Cixerri. Rendiconti dell’Associazione
Mineralogica Sarda 84: 43–65.
Amiranashvili NG. 2000. Differences in shell morphology of Testudo graeca and
Testudo hermanni, based on material from Bulgaria. Amphibia-Reptilia.
21:67–81. doi:10.1163/156853800507282.
Angelini G. 1899. Notizie de osservazioni intorno alla naturalizzazione della
Testudo nemoralis ‘Aldrov. Sardegna Bollettino della Società Romana per gli
studi zoologici. 8:50–52.
Angelone C, Moncunill-Solé B, Kotsakis T. 2020. Fossil Lagomorpha
(Mammalia) of Italy: systematics and biochronology. Rivista Italiana di
Paleontologia e Stratigrafia. 126:157–187. doi:10.13130/2039-4942/13014.
Azzaroli A. 1946. La scimmia fossile della Sardegna. Rivista di Scienze
Preistoriche. 1:68–76.
Baldino B. 2000. Resti faunistici dal Convento basso medievale di Saccargia
(Sassari). Italy: Atti 3°Convegno Nazionale di Archeozoologia (Siracusa); p.
501–505.
Barbieri F, Cherchi A. 1980. Excursion sur le Mésozoique et le Tertiaire de la
Sardaigne – livret-guide. C.i.e.,s.m C.N.R.; P.F. Geodinamica. 345:1–127.
Barca S, Melis E, Annino E, Cincotti F, Ulzega A, Orrù P, Pintus C. 2005. Note
Illustrative della Carta Geologica d’Italia alla scala 1:50.000. In: Foglio 557
“Cagliari”. Servizio Geologico d’Italia, Regione Autonoma della Sardegna. S.E
L.CA, Firenze.
Bate DMA. 1945. The Pleistocene mole of Sardinia. Ann Mag Natl Hist.
12:448–461. doi:10.1080/00222934508654743.
Bergounioux F-M. 1935. Contribution à l’étude paléontologique des chéloniens:
chéloniens fossiles du bassin d’Aquitaine. Mémoires de la Société Géologique
de France (nouvelle série). 25:1–215.
Bosco C. 1902. Il Lophiodon sardus (n. sp.) delle ligniti di Terras de Collu
(Sardegna). Rendiconti della Reale Accademia Nazionale dei Lincei.
11:178–182.
Brandy LD. 1978. 1905. Données nouvelles sur l’évolution du rongeur
endémique fossile corso-sarde Rhagamys. Major F Mammalia, Rodentia:
Bulletin de la Société Géologique de France; Vol. 20, p. 831–835.
Bringsøe H, Buskirk JR, Willemsen RE. 2001. Testudo marginata Schoepff, 1792-
Breitrandschildkröte. In: Fritz U, editor. Handbuch der Reptilien und
Amphibien Europas. Band 3/IIIA. Wiebelsheim: Aula-Verlag; p. 291–334.
Broin de F. 1977. Contribution à l’étude des Chéloniens: chéloniens continen­
taux du Crétacé et du Tertiaire de France. Mémoires du Muséum National
d’Histoire Naturelle, Série C. 38:1–366.
Caloi L, Kotsakis T, Palombo MR, Petronio C. 1980. Il giacimento a vertebrati
del Pleistocene superiore di San Giovanni in Sinis (Sardegna occidentale).
Atti della Accademia Nazionale dei Lincei Classe di Scienze Fisiche,
Matematiche e Naturali – Rendiconti. 69:185–197.
Capellini G. 1890a. Sul coccodrilliano gavialoide (Tomistoma calaritanus) sco­
perto nella collina di Cagliari nel 1868. Rendiconti della Reale Accademia dei
Lincei. 4:149–151.
Capellini G. 1890b. Sul coccodrilliano garialoide (Tomistoma calaritanus) sco­
perto nella collina di Cagliari nel MDCCCLXVIII. Atti della Reale Accademia
Lincei, Memorie della Classe di Scienze Fisiche. Matematiche e Naturali.
6:507–533.
Cappetta H, Thaler L. 1974. Présence de poissons Characidae, caractéristiques de
l’Éocène inferieur éuropeen, dans la formation lignitifere en Sardaigne.
Paleogeografia del Terziario sardo nell’ambito del Mediterraneo occidentale,
Rendiconti del Seminario della Facoltà di Scienze dell’Università di Cagliari.
41:69–71.
Carboni S, Lecca L. 1985. Osservazioni sul Pleistocene superiore della penisola
del Sinis (Sardegna occidentale). Bollettino della Società Geologica Italiana.
104:459–477.
Carboni S, Lecca L. 1995. Le Pliocène de Capo Mannu (Sardaigne occidentale):
transition marin littoral-continental dunaire. Comptes Rendus de l’Académie
des Sciences. 320:1203–1210.
Carmignani L, Oggiano G, Funedda A, Conti P, Pasci S, Barca S. 2008. Carta
geologica della Sardegna. Scala. 1:250.000. Servizio Geologico d’Italia.
Casanovas-Vilar I, van Dam JA, Trebini L, Rook L. 2011. The rodents from the
Late Miocene Oreopithecus-bearing site of Fiume Santo (Sardinia, Italy).
Geobios. 44:173–187. doi:10.1016/j.geobios.2010.08.002.
Cherchi A. 1974. Appunti biostratigrafici sul Miocene della Sardegna (Italia),
actes V congrès du Néogene Méditerranée. Mémoire du Bureau de
Recherches Géologiques et Minières 78: 433–445.
Cherchi A. 1979. Microfaune aptiano-(?) albiane dei ciottoli urgoniani della
Formazione del Cixerri (Sardegna SW) e loro interesse paleogeografico.
Rivista Italiana Di Paleontologia E Stratigrafia. 85:353–410.
Chesi F, Delfino M, Abbazzi L, Carboni S, Lecca L, Rook L. 2007a. New fossil
vertebrate remains from San Giovanni di Sinis (Late Pleistocene, Sardinia):
the last Mauremys (Reptilia, Testudines) in the central Mediterranean. Rivista
Italiana di Paleontologia e Stratigrafia. 113:287–297. doi:10.13130/2039-
4942/5875.
Chesi F, Delfino M, Varola A, Rook L. 2007b. Fossil sea turtles (Chelonii,
Dermochelyidae and Cheloniidae) from the Miocene of Pietra Leccese (late
Burdigalian-early Messinian), Southern Italy. Geodiversitas. 29:321–333.
Chesi F 2008. Il registro fossile italiano dei cheloni. Università di Firenze,
Florence, Unpublished PhD dissertation. pp. 1–178.
Chesi F, Delfino M, Pillola GL, Rook L, Villani M. 2008. A Pleistocene European
pond turtle from Sardinia. In: Corti C, editor. Herpetologia Sardiniae. Latina:
Edizioni Belvedere; p. 138–141.
Chesi F, Delfino M, Rook L. 2009. Late Miocene Mauremys (Testudines,
Geoemydidae) from Tuscany (Italy): evidence of terrapin persistence after
a mammal turnover. J Paleontol. 83:379–388. doi:10.1666/08-134.1.
Cita MB, Abbate E, Aldighieri B, Balini M, Conti MA, Farloni P, Germani D,
Groppelli G, Manetti P, Petti FM. 2007. Catalogo delle formazioni - Unità
tradizionali, Carta Geologica d’Italia - 1:50.000. In: Quaderni serie. III, 7 (VI).
Servizio Geologico d’Italia, Roma: Dipartimento Difesa del suolo; p. 318. APAT -.
Cocozza T, Decandia FA, Gandin A. 1986. Studio geologico stratigrafico
e paleogeografico del bacino carbonifero del Sulcis, nel programma di
ricerche minerarie di base. In: Convenzione Società Carbosulcis
e Università di Siena, Relazione inedita. Siena; p. 88.
Collareta A, Casati S, Zuffi MAL, Di Cencio A. 2020. First authentic record of the
freshwater turtle Mauremys from the Upper Pliocene of Italy, with a new
occurrence of the rarely reported ichnotaxon Thatchtelithichnus holmani.
Carnets de Géologie. 20:301–313. doi:10.2110/carnets.2020.2016.
Colombero S, Alba DM, D’Amico C, Delfino M, Esu D, Giuntelli P,
Harzhauser M, Mazza PPA, Mosca M, Neubauer TA, et al. 2017. Late
Messinian mollusks and vertebrates from Moncucco Torinese,
north-western Italy. Paleoecol Paleoclimatol Implications Palaeontol
Electronica. 20(1.10A):1–66.
Comaschi Caria I. 1959. Nuovi resti di cheloni nel Miocene della Sardegna.
Bollettino della Società Geologica Italiana. 78:37–44.
Corsini-Foka M, Masseti M. 2008. On the oldest record of the Nile soft-shelled
turtle, Trionyx triunguis (Forskål, 1775), in the Eastern Aegean islands
(Greece). Zool Middle East. 43:108–110. doi:10.1080/09397140.2008.
10638276.
Costamagna LG, Schäfer A. 2013. The Cixerri Fm (Middle Eocene-Early
Oligocene): analysis of a “Pyrenean” continental molassic system in southern
Sardinia. J Mediterr Earth Sci Spec. 41–44.
De Bruijn H, Rümke CG 1974. On a peculiar mammalian association from the
Miocene of Oschiri (Sardinia). Proceedings of the Koninklijke Nederlandse
Akademie Van Wetenschappen Series B. 77:46–79.
Del Vecchio C. 1921. Su alcuni denti di Tomistoma (Crocodilia) dell’Oligocene
di Visone presso Acqui. Atti della Società Italiana di Scienze Naturali.
60:419–431.
Delfino M, Bailon S. 2000. Early Pleistocene herpetofauna from Cava Dell’ Erba
and Cava Pirro (Apulia, Southern Italy). Herpetol J. 10:95–110.

Delfino M 2002. Erpetofaune italiane del Neogene e del Quaternario. PhD thesis
in Palaeontology, Università degli Studi di Modena e Reggio Emilia, Modena.
pp. 1–382.
Delfino M, Kotsakis T, Arca M, Tuveri C, Pitruzzella G, Rook L. 2008.
Agamid lizards from the Plio-Pleistocene of Sardinia (Italy) and an
overview of the European fossil record of the family. Geodiversitas.
30:641–656.
Delfino M, Pitruzzella G, Bailon S. 2011. The Late Pliocene amphibians and
reptiles from “Capo Mannu D1 Local Fauna” (Mandriola, Sardinia, Italy).
Geodiversitas. 33:357–382. doi:10.5252/g2011n2a10.
Delfino M, Luján ÀH, Carmona R, Alba DM. 2012. Revision of the extinct
Pleistocene tortoise Testudo lunellensis Almera and Bofill, 1903 from Cova de
Gràcia (Barcelona, Spain). Amphibia-Reptilia. 33(2):215–225. doi:10.1163/
156853812X636466.
Delfino M, Zoboli D, Carnevale G, Pillola GL. 2014. The rediscovered holotype
of Palaeopython sardus Portis, 1901 from the Miocene of Sardinia belongs to
a fish, not a snake. Bollettino della Società Paleontologica Italiana. 53:89–92.
doi:10.4435/BSPI.2014.09.
Delussu F. 2000. Lo stato attuale degli studi sulle faune oloceniche della Sardegna
centrosettentrionale. In: Giacobini G, Riede A, Tagliacozzo A, editors. Atti 2°
Convegno Nazionale Archeozoologia (Asti, 1997). Forlì: ABACO Edizioni; p.
183–192.
Depéret C, and Donnezan A. 1890–1897. Classe des reptiles. In: Depéret C,
editor. Les animaux Pliocènes du Roussillon. Vol. 3, Paris: Mémoires de la
Société Géologique de France, Paléontologie; p. 140–194.
Depéret, C. 1897. Etude de quelques gisements nouveaux de Vertébrés
pléistocènes de l’île de Corse. Annales de la Société Linnéenne de Lyon 44:
111–128.
Esu D, Kotsakis T. 1980. Presenza di Hypnomys Bate (Gliridae, Rodentia) nel
Villafranchiano di Nuraghe Su Casteddu (Nuoro, Sardegna). Atti della
Accademia Nazionale dei Lincei, Classe di Scienze Fisiche, Matematiche
e Naturali, Rendiconti. 8:123–127.
Esu D, Kotsakis T. 1983. Les vertébrés et les mollusques continentaux du
Tertiaire de la Sardaigne: paléobiogéographie et biostratigraphie. Geologica
Romana. 22:177–206.
Fanni S, Murru M, Salvadori I, Sarria E. 1982. Nuovi dati strutturali sul bacino
del Sulcis. L’Industria Mineraria. 4:25–31.
Fitzinger L. 1843. Systema Reptilium, fasciculus primus, Amblyglossae. Wien:
Braumüller et Seidel; p. 1–106.
Forskål P. 1775. Descriptiones Animalium. Avium, Amphibiorum, Piscium,
Insectorum, Vermium. Quae in Itinere Oriental Observavit. Haunia,
Denmark: Mölleri; p. 1–12.
Fritz U. 1995. Zur innerartlichen Variabilität von Emys orbicularis (Linnaeus,
1758). 5 Taxonomie in Mittel-Westeuropa, auf Korsika, Sardinien, der
Apenninen-Halbinsel und Sizilien und Unterartengruppen von E. orbicularis
Zoologische Abhandlungen. 48:185–242.
Fritz U, Fattizzo T, Guicking D, Tripepi S, Pennisi MG, Lenk P, Joger U,
Wink M. 2005a. A new cryptic species of pond turtle from southern Italy,
the hottest spot in the range of the genus Emys. Zool Scr. 34:351–371. doi:10.
1111/j.1463-6409.2005.00188.x.
Fritz U, Siroky P, Kami H, Wink M. 2005b. Environmentally caused dwarf­
ism or a valid species – is Testudo weissingeri Bour, 1996 a distinct
evolutionary lineage? New evidence from mitochondrial and nuclear
genomic markers. Mol Phylogenet Evol. 37:389–401. doi:10.1016/j.
ympev.2005.03.007.
Fucini A. 1912. Trionyx pliocenicus Law. Palaeontographia italica. 28:1–28.
Funedda A, Carmignani L, Pasci S, Patta ED, Uras V, Conti P, Sale V. 2009. Note
Illustrative della Carta Geologica d’Italia alla scala 1:50.000. In: Foglio 556
“Assemini”. Servizio Geologico d’Italia, Regione Autonoma della. Sardegna,
S.E L.CA, Firenze.
Gaffney ES. 1996. The postcranial morphology of Meiolania platyceps and
a review of the meiolaniidae. Bull Am Mus Nat Hist. 229:1–166.
Gandolfi R, Porcu A. 1967. Contributo alla conoscenza delle microfacies mio­
ceniche delle colline di Cagliari (Sardegna). Rivista Italiana di Paleontologiae
Stratigrafia. 73:313–348.
Gardner JD, Russell AP. 1994. Carapacial variation among soft-shelled turtles
(Testudines: Trionychidae), and its relevance to taxonomic and systematic
studies of fossil taxa. Neues Jahrbuch für Geologie und Paläontologie,
Abhandlungen. 193:209–244.
Gattacceca J, Deino AL, Rizzo R, Jones B, Henry B, Beaudoin B, Vadeboin F.
2007. Miocene rotation of Sardinia: new paleomagnetic and geochronological
constraints and geodynamic implications. Earth Planet Sci Lett. 258:359–377.
doi:10.1016/j.epsl.2007.02.003.
Gennari P. 1868. Di un coccodrillo fossile nel terreno pliocenico di Cagliari. Atti
dell’Accademia dei Fisiocritici di Siena. 2:127–129.
HISTORICAL BIOLOGY 27
Geoffroy Saint-Hilaire EF. 1809. Mémoire sur les tortues molles, nouveau genre
sous le nom de Trionyx, et sur la formation des carapaces. Annales du
Muséum d’Histoire Naturelle. 14:1–20.
Georgalis GL, Kear BP. 2013. The fossil turtles of Greece: an overview of
taxonomy and distribution. Geobios. 46:299–311. doi:10.1016/j.geobios.
2013.05.001.
Georgalis GL, Velitzelos E, Velitzelos D, Kear BP. 2013. Nostimochelone lampra
gen. et sp. nov., an enigmatic new podocnemidoidean turtle from the Lower
Miocene of Northern Greece. In: Brinkman D, Holroyd P, Gardner J, editors.
- Morphology and evolution of turtles: papers in honor of Eugene S Gaffney.
Vol. 3, Pleurodires. Dordrecht: Springer; p. 277–287. doi:10.1007/978-94-
007-4309-0_17.
Georgalis GL, Villa A, Vlachos E, Delfino M. 2016. Fossil amphibians and
reptiles from Plakias, Crete: a glimpse into the earliest late Miocene herpe­
tofaunas of Southeastern Europe. Geobios. 49:433–444. doi:10.1016/j.geo
bios.2016.09.004.
Georgalis GL, Joyce WG. 2017. A review of the fossil record of Old World turtles
of the clade Pan-Trionychidae. Bull Peabody Mus Nat Hist. 58:115–208.
doi:10.3374/014.058.0106.
Georgalis GL, Zoboli D, Pillola GL, Delfino M. 2017. A revision of the trionychid
turtle Procyclanorbis sardus Portis, 1901 from the late Miocene of Sardinia
(Italy). Annales de Paléontologie. 103:127–134. doi:10.1016/j.annpal.2017.04.
002.
Georgalis GL, Arca M, Rook L, Tuveri C, Delfino M. 2019. A new colubroid
snake (Serpentes) from the early Pleistocene of Sardinia, Italy. Bollettino
della Società Paleontologica Italiana. 58:277–294. doi:10.4435/BSPI.2019.
19.
Georgalis GL, Smith KT. 2020. Constrictores Oppel, 1811 - the available name
for the taxonomic group uniting boas and pythons. Vertebr Zool.
70:291–304. doi:10.26049/VZ70-3-2020-03.
Georgalis GL, Insacco G, Rook L, Spadola F, Delfino M. 2020a. Turtle remains
from the late Miocene of the Cessaniti area, southern Italy-insights for
a probable Tortonian chelonian dispersal from Europe to Africa. Swiss
J Palaeontol. 139:1. doi:10.1186/s13358-020-00202-y.
Georgalis GL, Zoboli D, Pérez-Garcìa A, Pillola GL, Delfino M. 2020b. The
occurence of Eocenochelus (Testudines, Pleurodira) from Sardinia supports
palaeogeographic reconstruction of the proximity of the island to continental
Western Europe during the Eocene. Rivista Italiana di Paleontologia
e Stratigrafia. 126(3):833–846. doi:10.13130/2039-4942/14443.
Georgalis GL, Delfino M. 2021. The Scontrone turtles – a new insular testudi­
noid fauna from the late Miocene of the Central Mediterranean. Geobios.
68:71–81. doi:10.1016/j.geobios.2021.05.001.
Georgalis GL, Macaluso L, Delfino M. 2021. A review of the fossil record of
Afro-Arabian turtles of the clade Testudinoidea. Bull Peabody Mus Nat Hist.
62:43–78. doi:10.3374/014.062.0103.
Gliozzi E, Malatesta A. 1980. The Quaternary goat of Capo Figari (Northeastern
Sardinia). Geologica Romana. 19:295–347.
Gmelin, JF. 1789. Caroli a Linné. Systema naturae per regna tria natural,
secundum classes, ordines, genera, species, cum characteribus differentilis,
synonymis, locis. Tomus I, Editio decima tertia, aucta, reformata. Pars III.
Amphibia et Pisces. Georg. Emanuel Beer, Lipsiae.
Gray, JE. 1864. Revision of the species of Trionychidae found in Asia and Africa,
with the description of some new species. Proceedings of the Zoological
Society of London 1864. London. pp. 76–98.
Gray JE. 1872. Appendix to the Catalogue of Shield Reptiles in the Collection of
the British Museum. Part I. Testudinata (Tortoises). London: Taylor and
Francis; p. 1–28.
Hensel, RF. 1856. Beiträge zur Kenntnis fossiler Säugetiere. II: Ueberreste von Mus
in der Breccie von Cagliari Zeitschrift der Deutschen Geologischen Gesellshaft.
8: 281–289.
Hervet S. 2000. Tortues du Quaternaire de France: critères de détermination,
répartitions chronologique et géographique. Mésogée. 58:3–47.
Joyce WG, Bell CJ. 2004. A review of the comparative morphology of extant
testudinoid turtles (Reptilia: Testudines). Asiatic Herpetol Res.
10:53–109.
Joyce WG. 2007. Phylogenetic relationships of Mesozoic turtles. Bull Peabody
Mus Nat Hist. 48(1):3–102. doi:10.3374/0079-032X.2007 48
Joyce WG, Anquetin J, Cadena E-A, Claude J, Danilov IG, Evers SW,
Ferreira GS, Gentry AD, Georgalis GL, Lyson TR, et al. 2021.
A nomenclature for fossil and living turtles using phylogenetically
defined clade names. Swiss J Palaeontol. 140:5. doi:10.1186/s13358-
020-00211-x.
Klembara J. 1979. Neue Funde der Gattungen Ophisaurus und Anguis
(Squamata, Reptilia) aus dem Untermiozän Westböhmens (CSSR). Vestník
Ústredního ústavu geologického. 54:163–169.
28 D. ZOBOLI ET AL.
Klembara J, Rummel M. 2018. New material of Ophisaurus, Anguis and
Pseudopus (Squamata, Anguidae, Anguinae) from the Miocene of the
Czech Republic and Germany and systematic revision and palaeobiogeogra­
phy of the Cenozoic Anguinae. Geol Mag. 155:20–44. doi:10.1017/
S0016756816000753.
Kordikova EG. 2002. Heterochrony in the evolution of the shell of Chelonia.
Part 1: terminology, Cheloniidae, Dermochelyidae, Trionychidae,
Cyclanorbidae and Carettochelyidae. Neues Jahrbuch für Geologie und
Paläontologie, Abhandlungen. 226:343–417. doi:10.1127/njgpa/226/2002/
343.
Kotsakis T. 1980. I resti di Anfibi e rettili Pleistocenici della Grotta di Dragonara
(Capo Caccia, Sardegna). Geologica Romana. 19:85–90.
Kotsakis T. 1981. Gli anfibi e i rettili dei Pleistocene del Lazio (Italia Centrale).
Geologica Romana. 20:57–67.
Kotsakis T. 1983. I Rettili olocenici della grotta Su Guanu (Nuoro, Sardegna).
Bollettino della Società Sarda di Scienze Naturali. 21:121–128.
Kotsakis T. 1985. Les Trionychidae fossils dell’Italie. Bollettino della Società
Paleontologica Italiana. 24:161–168.
Kotsakis T, Barisone G, Rook L. 1997. Mammalian biochronology in an insular
domain: the Italian Tertiary faunas. Mémoires et Travaux de l’Institut de
Montpellier. 21:431–441.
Kotsakis T, Murru M, and Palombo MR. 2008a. Terras de Collu. In:
Palombo MR, Pillola GL, Kotsakis T, editors. Euromam 2008 - Fossil
Mammalian Biotas of Sardinia Fieldtrip Guide-Book (Cagliari). p. 1–98.
Kotsakis T, Palombo MR, Angelone C, Melis RT, and Fanelli F. 2008b.
Mandriola - Capo Mannu. In: Palombo MR, Pillola GL, Kotsakis T, editors.
Euromam 2008 - Fossil Mammalian Biotas of Sardinia Fieldtrip Guide-
Book (Cagliari). p. 1–98.
Lapparent de Broin de F. 2001. The European turtle fauna from the Triassic to
the Present. Dumerilia. 4:155–216.
Lapparent de Broin de F. 2002. A giant tortoise from the Late Pliocene of Lesvos
Island (Greece) and its possible relationships. Annales Géologiques des Pays
Helléniques. 39:99–130.
Lecca L, Lonis R, Luxoro S, Melis E, Secchi F, Brotzu P. 1997. Oligo-Miocene
volcanic sequences and rifting stages in Sardinia: a review. Periodico di
Mineralogia. 66:7–61.
Lecca L, Carboni S. 2007. The Tyrrhenian section of San Giovanni di Sinis
(Sardinia): stratigraphic record, of an irregular single high stand. Rivista
Italiana di Paleontologia e Stratigrafia. 113(3):509–523. doi:10.13130/2039-
4942/5889.
Leith Adams A. 1877. On gigantic land-tortoises and a small freshwater species
from the ossiferous caverns of Malta, together with a list of their fossil fauna;
and a note on chelonian remains from the rock cavities of Gibraltar. Q J Geol
Soc London. 33:177–191. doi:10.1144/GSL.JGS.1877.033.01-04.11.
Levine M. 1983. La fauna di Filiestru (trincea D). In: Trump DH, editor. La
grotta di Filiestru a Bonuighinu, Mara (SS): quaderni 13, Soprintendenza ai
Beni Archeologici delle province di Sassari e Nuoro 13 (Sassari): 111–131.
López Martínez N, Thaler L. 1975. Biogéographie, évolution et compléments à la
systématique du groupe d’Ochotonides Piezodus-Prolagus (Mammalia,
Lagomorpha). Bulletin de la Société Géologique de France. 17:850–866.
doi:10.2113/gssgfbull.S7-XVII.5.850.
Loveridge A, Williams EE. 1957. Revision of the African tortoises and turtles of
the suborder Cryptodira. Bull Mus Comp Zool. 115:163–557.
Luján ÀH, Alba D, Fortuny J, Carmona R, Delfino M. 2014. First cranial remains
of Cheirogaster richardi (Testudines: Testudinidae) from the Late Miocene of
Ecoparc de Can Mata (Vallés-Penedés Basin, NE Iberian Peninsula): taxo­
nomic and phylogenetic implications. J Syst Palaeontol. 12:833–864. doi:10.
1080/14772019.2013.863231.
Luján ÀH, Delfino M, Robles JM, Alba DM. 2016. The Miocene tortoise Testudo
catalaunica Bataller, 1926, and a revised phylogeny of extinct species of genus
Testudo (Testudines: Testudinidae). Zool J Linn Soc. 178(2):312–342. doi:10.
1111/zoj.12414.
Luján ÀH, Alcover JA, Ivanov M, Torres E, Alba DM. 2017. Revisión
taxonómica de “Testudo” gymnesica Bate, 1914 (Testudines, Testudinidae)
a partir de la descripción del material tipo de Menorca (Islas Baleares).
Spanish J Palaeontol. 32:261–278. doi:10.7203/sjp.32.2.17043.
Lydekker RA. 1891a. On a new species of Trionyx from the Miocene of Malta
and a chelonian scapula from the London Clay. Q J Geol Soc London.
47:37–40. doi:10.1144/GSL.JGS.1891.047.01-04.05.
Lydekker RA 1891b. On Pleistocene Bird-remains from the Sardinian and Corsican
Islands. Proceedings of the Zoological Society of London. 3:467–477.
Major CIF. 1882. L’origine della fauna delle nostre isole. Atti della Società
Toscana di Scienze Naturali, Processi Verbali. 3:36–42 and 113–133.
Major CIF 1901. Exhibition of, and remaks upon, the skull of a new fossil
mammal (Enhydrictis galictoides) from Sardinia. Proceedings of Zoological
Society of London London. pp. 625–628.
Major CIF. 1905. Rodents from the Pleistocene of the Western Mediterranean
region. Geol Mag. 2:501–506. doi:10.1017/S0016756800128493.
Massari F, Dieni L. 1973. La formazione fluvio-lacustre di Nuraghe Casteddu ed
i suoi rapporti con i basalti di Orosei-Dorgali (Sardegna). Memorie della
Società Geologica Italiana. 12:377–410.
Maxia C. 1959. Malacofauna oligotipica di età paleogenica della Valle del Cixerri
(Iglesiente). Pubblico Istituto Geologico Paleontologico Università di Roma.
95:1–18.
Mayer R. 1992. Europäische landschildkröten. Kempten, Allgäu: Leben–
Haltung–Zucht; p. 1–127.
Mazzei R, Oggiano G. 1990. Messa in evidenza di due cicli sedimentari nel
Miocene dell’area di Florinas (Sardegna settentrionale). Atti della Società
Toscana di Scienze Naturali, Memorie. 97:119–147.
Mennecart B, Zoboli D, Costeur L, Pillola GL. 2017. Reassessment of the
ruminant from Sardara, the last non-insular mammal from Sardinia (Italy).
Neues Jahrbuchfür Geologie und Paläontologie. 286:97–104. doi:10.1127/
njgpa/2017/0688.
Mennecart B, Zoboli D, Costeur L, Pillola GL. 2019. On the systematic position
of the oldest insular ruminant Sardomeryx oschiriensis (Mammalia,
Ruminantia) and the early evolution of the Giraffomorpha. J Syst
Palaeontol. 17(8):691–704. doi:10.1080/14772019.2018.1472145.
Meyer von H. 1843. Mittheilungen an Prof. Bronn Gerichtet Neues Jahrbuch Für
Mineralogie, Geognosie, Geologie Und Petrefaktenkunden 1843 . 698–704.
Meylan PA, Weig BS, Wood RC. 1990. Fossil soft shelled turtles (Family
Trionychidae) of the Lake Turkana Basin, Africa. Copeia. 1990:508–528.
doi:10.2307/1446355.
Młynarski M. 1976. Testudines, Part 7 In: Fischer Verlag, G. Handbook of
Palaeoherpetology. Stuttgart, New York: Gustav Fischer Verlag; p. 1–129.
Nicholl CC, Rio JP, Mannion PD, Delfino M. 2021. A re-examination of the anatomy
and systematics of the tomistomine crocodylians from the Miocene of Italy and
Malta. J Syst Palaeontol. 18:1853–1889. doi:10.1080/14772019.2020.1855603.
Oudet J, Münch PH, Verati C, Ferrandini M, Melinte-Dobrinescu M,
Gattacecca J, Cornée JJ, Oggiano G, Quillévéré F, Borgomano J, et al. 2010.
Integrated chronostratigraphy of an intra-arc basin: 40Ar/39Ar datings,
micropalaeontology and magnetostratigraphy of the early Miocene
Castelsardo basin (northern Sardinia, Italy). Palaeogeogr Palaeoclimatol
Palaeoecol. 295:293–306. doi:10.1016/j.palaeo.2010.06.007.
Palombo MR, Ibba A, and Fanelli F. 2008. Porto Conte Bay. In: Palombo MR,
Pillola GL, Kotsakis T, editors. Euromam 2008 - Fossil Mammalian Biotas of
Sardinia Fieldtrip Guide-Book (Cagliari). p. 1–98.
Palombo MR, Antonioli F, Lo Presti V, Mannino MA, Melis RT, Orrù P,
Stocchi P, Talamo S, Quarta G, Calcagnile L, et al. 2017. The late
Pleistocene to Holocene palaeogeographic evolution of the Porto Conte
area: clues for a better understanding of human colonization of Sardinia
and faunal dynamics during the last 30 ka. Quat Int. 439:117–140. doi:10.
1016/j.quaint.2016.06.014.
Palombo MR. 2018. Insular mammalian fauna dynamics and paleogeography:
a lesson from the Western Mediterranean islands. Integr Zool. 13:2–20.
doi:10.1111/1749-4877.12275.
Parham JF, Fastovsky DE. 1997. The phylogeny of cheloniid sea turtles revisited.
Chelonian Conserv Biol. 2:548–554.
Parham JF. 2005. A reassessment of the referral of sea turtle skulls to the
genus Osteopygis (Late Cretaceous, New Jersey, USA). J Vertebr
Paleontol. 25(1):71–77. doi:10.1671/0272-4634(2005)025[0071:
AROTRO]2.0.CO;2.
Pecorini G, Pomesano Cherchi A. 1969. Ricerche geologiche e biostratigrafiche
sul Campidano meridionale (Sardegna). Memorie della Società Geologica
Italiana. 8:421–451.
Pecorini G, Rage J-C, Thaler L. 1974. La Formation Continentale de Capo
Mannu, sa faune à Vertébrés pliocènes et la question du Messinies en
Sardaigne. Rendiconti del Seminario della Facoltà di Scienze dell’Università
di Cagliari. 33:305–320.
Pedall I, Fritz U, Stuckas H, Valdeón A, Wink M. 2011. Gene flow across
secondary contact zones of the Emys orbicularis complex in the Western
Mediterranean and evidence for extinction and re-introduction of pond
turtles on Corsica and Sardinia (Testudines: Emydidae). J Zool Syst Evol
Res. 49:44–57. doi:10.1111/j.1439-0469.2010.00572.x.
Perez M, Leblois R, Livoreil B, Bour R, Lambourdiere J, Samadi S,
Boisselier M-C. 2012. Effects of landscape features and demographic history
on the genetic structure of Testudo marginata populations in the southern
Peloponnese and Sardinia. Biol J Linn Soc. 105:591–606. doi:10.1111/j.1095-
8312.2011.01805.x.
Perez M, Livoreil B, Mantovani S, Boisselier M-C, Crestanello B, Abdelkrim J,
Bonillo C, Goutner V, Lamb Ourdière J, Pierpaoli M, et al. 2014. Genetic
variation and population structure in the Endangered Hermann’s Tortoise:
the roles of geography and human-mediated processes. J Heredity.
105:70–81. doi:10.1093/jhered/est071.

Pérez-García A, Vlachos E. 2014. New generic proposal for the European
Neogene large testudinids (Cryptodira) and the first phylogenetic hypothesis
for the medium and large representatives of the European Cenozoic record.
Zool J Linn Soc. 172:653–719. doi:10.1111/zoj.12183.
Pérez-García A, Lapparent de Broin F, Murelaga X. 2017a. The Erymnochelys
group of turtles (Pleurodira, Podocnemididae) in the Eocene of Europe: new
taxa and paleobiogeographical implications. Palaeontol Electronica. 20
(1.14A):1–28.
Pérez-García A, Vlachos E, Arribas A. 2017b. The last giant continental tortoise
of Europe: a survivor in the Spanish Pleistocene site of Fonelas P-1.
Palaeogeogr Palaeoclimatol Palaeoecol. 470:30–39. doi:10.1016/j.palaeo.
2017.01.011.
Pérez-García A, Díaz-Berenguer E, Badiola A, Canudo JI. 2019. An unexpected
finding: identification of the first complete shell of the Franco-Belgian middle
Eocene littoral pleurodiran turtle Eocenochelus eremberti in Spain. Hist Biol.
33:527–533. doi:10.1080/08912963.2019.1644330.
Pomesano Cherchi A 1971. Microfaune plantoniche di alcune serie mioceniche
del Logudoro (Sardegna). In: Proceeding of the 2nd Planktonic Conference
Rome. pp. 1003–1066.
Porcu A. 1983. Geologia del Graben di Ottana (Sardegna centrale). Rendiconti
del Seminario della Facoltà di Scienze dell’Università di Cagliari. 53:151–169.
Portis A. 1893. Contribuzioni alla storia fisica del bacino di Roma e studii sopra
l’estenzione da darsi al Pliocene superiore. Roma L: Roux; p. 1–513.
Portis A. 1901a. Il Palaeopython sardus Port. Nuovo pitonide del Miocene medio
della Sardegna. Bollettino della Società Geologica Italiana. 20:247–253.
Portis A. 1901b. Il Procyclanorbis sardus Port., nuovo trionychide fossile della
Sardegna. Bollettino della Società Geologica Italiana. 20:51–79.
Pritchard PCH. 1988. A survey of neural bone variation among recent chelonian
species, with functional interpretations. Acta Zool Cracoviensia. 31:625–686.
Rhodin AGJ, Iverson J, Bour R, Fritz U, Georges A, Shaffer BH, van Dijk PP. 2021.
Turtles of the world: annotated checklist and atlas of taxonomy, synonymy, dis­
tribution, and conservation status (9th Ed.). Chelonian Res Monogr. 8:1–472.
Ristori G. 1895a. Cheloniani fossili di Montebamboli e Casteani. Memoria
paleontologica del prof. Giuseppe Ristori. Con appendice sui Cheloniani
fossili del Casino (Siena). Pubblicazioni del Reale Istituto di Studi superiori
in Firenze, ezione di scienze fisiche e naturale. 21:1–104.
Ristori G. 1895b. Di un nuovo Chelonio fossile del Miocene dell’Isola di Malta.
Atti Della Società Toscana Di Scienze Naturali, Memorie. 14:3–17.
Roček Z. 1984. Lizards (Reptilia, Sauria) from the lower Miocene locality
Dolnice (Bohemia, Czechoslovakia). Rozpravy Ceskoslovenské Akademie
Ved, Rada Matematickych a prírodních Ved. 94:3–69.
Rook L, Abbazzi A, Angelone C, Arca M, Barisone G, Bedetti C, Delfino M,
Kotsakis T, Marcolini F, Palombo MR, et al. 2003. Osservazioni preliminari
sui vertebrati fossili Plio-Pleistocenici del Monte Tuttavista (Orosei,
Sardegna). Int J Archaeol- Sardinia Corsica et Baleares Antiquae. 1:11–29.
Rook L, Bartolini Lucenti S, Tuveri C, Arca M. 2018. Mustelids (Carnivora,
Mammalia) from Monte Tuttavista fissure fillings (early and middle
Pleistocene; Orosei, Sardinia): taxonomy and evolution of the insular
Sardinian Galictini. Quat Sci Rev. 197:209–223. doi:10.1016/j.quascirev.
2018.08.022.
Sacco F. 1889. I Cheloni astiani del Piemonte. Memorie della Reale Accademia
delle Scienze di Torino. 39:427–461.
Sanchiz B. 1998. Salientia. In: Handbuch der Paläoherpetologie. Vol. 4.
München: Verlag Friedrich Pfeil; p. 1–275.
Sindaco R, Doria G, Razzetti E, Bernini F. 2006. Atlante degli Anfibi e dei Rettili
d’Italia/Atlas of Italian Amphibians and Reptiles. Societas Herpetologica
Italica, Edizioni Polistampa, Firenze. 1–789.
Sondaar PY, De Boer PL, Sanges M, Kotsakis T, Esu D. 1984. First report
on a Paleolithic culture in Sardinia. British Archaeol Rep Int Ser.
229:29–47.
Sowerbutts A. 2000. Sedimentation and volcanism linked to multiphase rifting
in an Oligo-Miocene intra-arc basin, Anglona Sardinia. Geol Mag.
137:395–418. doi:10.1017/S0016756800004246.
Spano S. 1985. Nuova segnalazione di resti di coccodrillo nel Miocene della
Sardegna. Bollettino della Società Sarda di Scienze Naturali. 24:1–12.
Speranza F, Villa IM, Sagnotti L, Florindo F, Cosentino D, Cipollari P, Mattei M.
2002. Age of the Corsica–Sardinia rotation and Liguro–Provençal Basin
spreading: new paleomagnetic and Ar/Ar evidence. Tectonophysics.
347:231–251. doi:10.1016/S0040-1951(02)00031-8.
Speybroeck J, Beukema W, Dufresnes C, Fritz U, Jablonski D, Lymberakis P,
Martínez-Solano I, Razzetti E, Vamberger M, Vences M, et al. 2020. Species
list of the European herpetofauna – 2020 update by the Taxonomic
Committee of the Societas Europaea Herpetologica. Amphibia-Reptilia. 41
(2):139–189. doi:10.1163/15685381-bja10010.
HISTORICAL BIOLOGY 29
Szalai T. 1931. Schildkrötenstudien. I. Testudo Schafferi nov. sp., eine
Riesenschildkröte aus dem Pliozän von Samos. Annalen des
Naturhistorischen Museums in Wien. 46:153–157.
Szyndlar Z. 1994. Oligocene snakes of southern Germany. J Vertebr Paleontol.
14:24–37. doi:10.1080/02724634.1994.10011536.
Taricco M. 1924. Il bacino lignitifero di Gonnesa (Provincia di Cagliari).
Bollettino del Regio Ufficio Geologico d’Italia. 49:1–14.
Taskavak E, Reimann MJ, Polder WN. 1999. First record of the Nile softshelled
Turtle, Trionyx triunguis, from Kos Island, Greece, with comments on its
occurrence in the eastern Mediterranean. Chelonian Conserv Biol.
3:510–512.
Tschopp E, Villa A, Camaiti M, Ferro L, Tuveri C, Rook L, Arca M,
Delfino M. 2018. The first fossils of Timon (Squamata: Lacertinae)
from Sardinia (Italy) and possible causes for its local extinction in the
Pleistocene. Zool J Linn Soc. 20:1–32. doi:10.1093/zoolinnean/zly003.
Valenti P, Vlachos E, Kehlmaier C, Fritz U, Georgalis GL, Luján AH, Miccichè R,
Sineo L, Delfino M. 2022. The last of the large-sized tortoises of the Mediterranean
islands. Zool J Linn Soc. doi:10.1093/zoolinnean/zlac044.
Vamberger M, Corti C, Stuckas H, Fritz U. 2011. Is the imperilled spur-thighed
tortoise (Testudo graeca) native in Sardinia? Implications from population
genetics and for conservation. Amphibia-Reptilia. 32:9–25. doi:10.1163/
017353710X541869.
Van der Made J. 1999. Biogeography and stratigraphy of the Mio-Pleistocene
mammals of Sardinia and the description of some fossils. In: Reumer JWF, de
Vos J, editors. Elephants have a snorkel! Papers in honour of PY Sondaar.
Deinsea. Vol. 7 (Rotterdam), p. 337–360.
Van der Made J. 2008. New endemic large mammals from the Lower Miocene of
Oschiri (Sardinia): observations on evolution in insular environment. Quat
Int. 182:116–134. doi:10.1016/j.quaint.2007.09.036.
Vardabasso S. 1962. Questioni paleogeografiche relative al Terziario antico della
Sardegna. Memorie della Società Geologica Italiana. 3:655–673.
Venczel M, Sanchiz B. 2006. Lower Miocene Amphibians and reptiles from
Oschiri (Sardinia, Italy). Hantkeniana. 5:72–75.
Villa A, Raineri G. 2015. The geologically youngest remains of Trachyaspis lardyi Meyer,
1843 (Testudines, Cheloniidae): a new specimen from the late Pliocene of the Stirone
River (Northern Italy). Bollettino della Società Paleontologica Italiana. 54:117–123.
doi:10.4435/BSPI.2015.07.
Villa M, Carnevale G, Pavia M, Rook L, Sami M, Szyndlar Z, Delfino M. 2021.
An overview on the late Miocene vertebrates from the fissure fillings of
Monticino Quarry (Brisighella, Italy), with new data on non-mammalian
taxa. Rivista Italiana di Paleontologia e Stratigrafia. 127:297–354. doi:10.
13130/2039-4942/15774.
Vitek NS, Joyce WG. 2015. A review of the fossil record of New World turtles of
the clade Pan-Trionychidae. Bull Peabody Mus Nat Hist. 56:185–244. doi:10.
3374/014.056.0204.
Vlachos E, Rabi M. 2018. Total evidence analysis and body size evolution of
extant and extinct tortoises (Testudines: Cryptodira: Pan-Testudinidae).
Cladistics. 34:652–683. doi:10.1111/cla.12227.
Vlachos E, Georgalis GL, Roussiakis S, Böhme M, Theodorou G. 2020a. The
Pikermian tortoises (Testudines, Testudinidae) from the late Miocene of the
South Balkans. J Vertebr Paleontol. 39:e1711520. doi:10.1080/02724634.2019.
1711520.
Vlachos E, Pérez-García A, Roussiakis S, Georgalis GL, Kear BP. 2020b. Late
Miocene tortoises from Samos, Greece: implications for European Neogene
testudinid systematics and distributions. J Vertebr Paleontol. 39:e1722950.
doi:10.1080/02724634.2019.1722950.
Wagner R. 1829. Beiträge zur Geschichte der fossilen Thiere. Leipzig: Isis von
Oken. Vol. 22, p.1132–1141.
Walker CA, Moody RTJ. 1974. A new trionychid turtle from the Lower Eocene
of Kent. Palaeontology. 17:901–907.
Wilkens B. 2003. Archeozoologia. Manuale per lo studio dei resti faunistici
dell’area mediterranea. Schio: Cd-rom.
Wyneken J. 2001. The anatomy of sea turtles. Miami: U.S. Department of
Commerce NOAA Technical Memorandum NMFS-SEFSC-470; p. 1–172.
Zammit Maempel G, de Bruijn H 1982. The Plio-Pleistocene Gliridae from the
Mediterranean islands reconsidered. Proceedings Koninklijke Nederlandse
Akademie van Wetenschappen. 85:113–128.
Zangerl R. 1969. The turtle shell. In: Gans C, D’a. Bellairs A, Parson TS, editors.
Biology of the Reptilia (Vol. I). London and New York: Morphology A,
Academic Press; p. 311–339.
Zenboudji S, Cheylan M, Arnal V, Bertolero A, Leblois R, Astruc G, Bertorelle G,
Pretus JL, Lo Valvo M, Sotgiu G, et al. 2016. Conservation of the endangered
Mediterranean tortoise Testudo hermanni hermanni: the contribution of
population genetics and historical demography. Biol Conserv. 195:279–291.
doi:10.1016/j.biocon.2016.01.007.
30 D. ZOBOLI ET AL.
Zoboli D, Pillola GL. 2016a. Quaternary mammal fauna from “Surconis”,
Bolotana (Sardinia, Italy). Bollettino della Società Paleontologica Italiana.
55:193–203. doi:10.4435/BSPI.2016.17.
Zoboli D, Pillola GL. 2016b. I rettili miocenici conservati nel Museo Sardo di
Geologia e Paleontologia Domenico Lovisato (Cagliari, Italia). Museologia
Scientifica – Nuova Serie. 10:81–87.
Zoboli D, Pillola GL. 2017. Early Miocene insular vertebrates from Laerru
(Sardinia, Italy): preliminary note. Rivista Italiana di Paleontologia
e Stratigrafia. 123(1):149–158. doi:10.13130/2039-4942/8073.
Zoboli D. 2017. I vertebrati quaternari sardi conservati nel Naturhistorisches
Museum di Basilea (Svizzera). Museologia Scientifica – Nuova Serie.
11:70–76.
Zoboli D, Sanciu L, Pillola GL, Delfino M. 2019. An overview of the crocodylian
fossil record from Sardinia (Italy). Annales de Paléontologie. 105:123–137.
doi:10.1016/j.annpal.2019.05.001.
Zoboli D, Pistis M, Afrasinei GM, Nonnoi G, Pillola GL. 2021. Crocodiles,
sharks and turtles: the urban geo‑palaeontological heritage of Cagliari
(Italy). Geoheritage. 13:52. doi:10.1007/s12371-021-00580-w.