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Mediterranean Quaternary interglacial molluscan assemblages:
Palaeobiogeographical and palaeoceanographical responses to climate change
Vittorio Garilli ⁎
APEMA-Research and Educational Service, Via Alla Falconara, 34, 90136 Palermo, Italy
abstractarticle info
Article history:
Received 16 May 2011
Received in revised form 15 September 2011
Accepted 26 September 2011
Available online 1 October 2011
Keywords:
Mollusca
Palaeobiogeography
Posidonia
Palaeoceanography
Pleistocene
Mediterranean
Atlantic
Africa
The analysis of several central to eastern Mediterranean shallow-water deposits, mostly related to Posidonia
oceanica palaeocommunities, sheds light on the presence of thermophilic molluscan assemblages throughout
the Mediterranean Pleistocene. The taxonomic composition of these assemblages is totally different from that
of the well-known Senegalese molluscan fauna and they play an important role in understanding some of the
hydroclimatic conditions during Quaternary interglacial episodes in the central-eastern Mediterranean basin.
Two of these assemblages from the Kyllini–Trypiti sequence (NW Peloponnesus) can be referred to the Upper
Pleistocene interglacial marine isotope (sub)stage (MIS) 5e and 5a, on the basis of U/Th dating of corals (Cla-
docora coespitosa). Other assemblages belong to Lower Pleistocene (mainly Emilian substage, possible MIS
37), late Lower (Sicilian substages)–early Middle Pleistocene and late Middle–Upper Pleistocene (MIS 7
and MIS 5e). As a whole, these assemblages consist of 15 species of palaeoclimatic and stratigraphic value.
Among these, Craspedochiton altavillensis (Polyplacophora), Jujubinus?bullula,Ersilia aliceae,Haedropleura
bucciniformis,Strioterebrum basteroti (Gastropoda), and Plicatula mytilina (Bivalvia) are (Euro)Mediterranean
strictly endemic extinct species, whereas Rissoina decussata,Niso terebellum,Kyllinia parentalis,Strioterebrum
grayi,Terebra corrugata,Terebra reticularis (Gastropoda), Anadara sp., Chama placentina, and Corbula revoluta
(Bivalvia) can be regarded as closely related to recent tropical West African taxa or as species living in trop-
ical waters. The latter stock is useful for reconstructing palaeoclimatic conditions, suggesting winter sea sur-
face temperatures 4–6 °C and 2–4 °C warmer than today in western Sicily and the easternmost part of the
Mediterranean Sea, respectively. In contrast with Pleistocene glacial settings, detected warm episodes are
characterized by a lower degree of seasonality, with a difference between summer and winter sea surface
temperatures of 6–7 °C vs. 10–11 °C recorded at present in the Mediterranean. The molluscan assemblages
also indicate that central to eastern Mediterranean salinity was lower than today, suggesting that a different
hydrological regime, triggered by increase in rainfall/runoff, was manifested during warmer Pleistocene pe-
riods in that area. The distribution of some of the most meaningful taxa studied here indicates that the Med-
iterranean Sea was divided into two palaeobioprovinces, a cooler western and a warmer eastern one, with a
boundary zone possibly located near the present Sicilian–Tunisian Strait.
The Anadara sp.–Jujubinus?bullula association is a meaningful ecobiostratigraphical marker for recognizing
warm Lower Pleistocene–Upper Pleistocene events. The youngest occurrence of this association has been
detected in MIS 5a, in the uppermost part of the Kyllini–Trypiti sequence. Craspedochiton altavillensis,J.?bul-
lula,Ersilia aliceae,Anadara sp., Chama placentina and Plicatula mytilina became extinct during the MIS 5a. Kyl-
linia parentalis and Terebra corrugata became extinct during MIS 5e. The disappearance of Niso terebellum
took place during the Emilian substage (Lower Pleistocene). The gastropod Nassarius musivus musivus, previ-
ously considered as becoming extinct at the top of the Gelasian stage, has been recorded in the eastern Med-
iterranean deposits younger than MIS 5a and older than MIS 3.
© 2011 Elsevier B.V. All rights reserved.
1. Introduction
Marine benthic molluscan associations have proved to be a power-
ful tool in recognizing palaeoenvironmental settings. In particular, the
duration of a certain range of temperature is the main factor in
controlling their latitudinal distribution and taxonomic diversity
(Hall, 1964; Monegatti and Raffi, 2001; Silva et al., 2010). Assuming
actualistic and uniformitarianism approaches, molluscan assemblages
can be used as reliable palaeoclimatic signals for the Quaternary.
These approaches were applied to molluscs in order to reconstruct
palaeoclimatic conditions, for example, of sea surface temperatures
(SSTs), deduced from palaeobiogeographical information (Raffiet al.,
1985; Monegatti and Raffi, 2001, 2007; Meco et al., 2002; Aguirre
et al., 2005; Bardají et al., 2009; Silva et al., 2010).
Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
⁎Tel.: +39 3291884289.
E-mail addresses: vittoriogarilli@apema.eu,vittoriogarilli@tiscali.it.
0031-0182/$ –see front matter © 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2011.09.012
Contents lists available at SciVerse ScienceDirect
Palaeogeography, Palaeoclimatology, Palaeoecology
journal homepage: www.elsevier.com/locate/palaeo
Author's personal copy
For the Mediterranean area, whereas many studies were dedicated
to molluscan biodiversity changes during Quaternary cold episodes
(e.g. Gignoux, 1913; Mars, 1963; Malatesta and Zarlenga, 1986; Raffi,
1986; Taviani et al., 1991; Di Geronimo et al., 2000), much less is
known about changing conditions throughout Quaternary Mediterra-
nean warm climatic settings, particularly for the Lower Pleistocene. In
fact, studies on Quaternary thermophilic assemblages appear consis-
tently limited to the late Middle–Upper Pleistocene interglacials, for
very few deposits of marine isotope stage (MIS) 7 (ca. 180–250 ka),
for MIS 9 (ca. 300–350 ka) or for MIS 11 (only at the Balearic Islands)
(ca. 420–360 ka) and, above all, for MIS 5e (in the Tyrrhenian stage;
ca. 135–116 ka) from the western part of the Mediterranean (Hearty
et al., 1986 with references; Hillaire-Marcel et al., 1996; Zazo et al.
2003a; Ferranti et al., 2006, with references; Bardají et al., 2009).
These are correlated with (warm) interglacial periods by global δ
18
O
stratigraphy (Gibbard and Kolfschoten, 2004). No consistent progress
in the study of warm phases/molluscs from Mediterranean Lower
Pleistocene has been obtained after the concise and pioneering
works of Ruggieri et al. (1982) and Ruggieri (1987a, 1987b). These au-
thors recognized Lower Pleistocene warm episodes by means of the
so-called “Pliocene resuscitates”(sensu Ruggieri, 1987a, 1987b), spe-
cies that originated in the Mio-Pliocene and re-immigrated into the
Mediterranean or survived the climatic deterioration at the Gela-
sian–Calabrian (sensu Gibbard et al., 2010) boundary.
The contribution of the present article mainly arises from the
study of Lower to Upper Pleistocene marine shallow-water molluscan
assemblages sampled in the south-central (Sicily, Italy) and eastern
(Peloponnesus, Greece) Mediterranean area. It includes a critical re-
view of other Mediterranean Quaternary central to eastern assem-
blages/deposits from bibliographic sources and museum collections
(Fig. 1). The majority of these molluscs are scarcely known or simply
have never been highlighted as a useful, reliable tool for reconstruct-
ing Pleistocene climatic settings.
The main aims of this article are: 1) to identify thermophilic ma-
rine molluscan species/assemblages from the Mediterranean Pleisto-
cene; 2) to focus on their biogeographical and biodiversity patterns
and to recognize their palaeoclimatic significance; 3) to delineate
their palaeoeocological and stratigraphical setting; 4) to compare
them with molluscs living along the West African coast; and 5) to at-
tempt a reconstruction of the central to eastern Mediterranean Pleis-
tocene SSTs and sea surface salinity (SSS) occurred during warm
episodes.
2. The Mediterranean Sea
2.1. Present-day
The Mediterranean (Fig. 1) is an almost landlocked basin with nat-
ural communications with the Atlantic Ocean, through the Straits of
Gibraltar, and the Black Sea, through the Bosporus, the Dardanelles,
and the Sea of Marmora. In 1869 the easternmost part of the basin
was modified by the cutting of the Suez Canal, allowing connection
with the Red Sea. This new route triggered the so-called Lessepsian
migration of several Red Sea–Indo-Pacific marine taxa into Mediterra-
nean (Barash and Danin, 1973; CIESM.ORG, 2002, with references).
The present-day Mediterranean hydroclimate is characterized by
an eastwards increase in monthly mean sea surface temperatures
(MMSSTs). Temperatures rise in August to about 23 °C in the Alboran
Sea, around the Straits of Gibraltar, 26 °C in western Sicily, and 28.5 °C
north of Cyprus (Figs. 2A and 3). Minimum temperatures occur in
February, reaching 17 °C in the south-eastern part, 15–16 °C in the
south-central part and in the vicinity of the Straits of Gibraltar, and
Quaternary recovery sites of the molluscan
assemblages in this study
Sampled for this study
Studied from bibliographic sources
Studied from Museum collections
Fig. 1. Location of the study area in the Mediterranean Sea. The black curves a–d indicate the western/eastern bioprovince boundaries following different authors.
From Bianchi (2007), with references.
99V. Garilli / Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
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10–13 °C in the north (see Figs. 2B and 3), with a trend throughout the
Mediterranean of surface isotherms matching that averaged over
1906–1995 (Bianchi, 2007, Fig. 2, with references). In the south-cen-
tral (western Sicily and north of Crete) and eastern parts of the basin
MMSSTs never go below 19–20 °C for at least six and eight months
of the year, respectively (Fig. 3). MMSSTs ≥24 °C occur for three and
five months of the year in the south-central (western Sicily) and the
eastern Mediterranean (Cyprus), respectively (Fig. 3). This is very like-
ly the reason that about 100 tropical Lessepsian molluscs are confined
to the easternmost coasts of the Mediterranean (CIESM.ORG, 2002),
whereas very few species maintain stable populations with difficulty
(Garilli and Caruso, 2004; Crocetta et al., 2009) north of the 15 °C
isotherm and never occur west of Sicily. The Mediterranean is
therefore generally subtropical, although under a process of tropicali-
zation, and has strong seasonality, especially in the central-eastern
portion, where the range between maximum (summer) and mini-
mum (winter) MMSSTs is of about 10–11 °C at present.
Annual salinity averaged over 10–100 m depth consistently varies
from west to east: 36 psu in the area of Gibraltar, 37–38 psu off the
western Sicilian coasts, 39 psu along the northwestern Peloponnesus
and 40 psu in the remaining eastern part of the Mediterranean
(NOAA WOA, 2001). No consistent differences are recorded between
rainy and dry seasons.
By means of biogeographical criteria, the Mediterranean has been
subdivided in 10–12 sectors (Bianchi, 2007), although the major dif-
ferentiation is that of a western and an eastern Mediterranean.
Opinions on the position of the western-eastern boundary are far
from perfectly concordant (see Fig. 1).
2.2. Brief Mio-Pleistocene history
After closing of the communication with the Indo-Pacific region,
the Mediterranean province represented a more-or-less independent
Atlantic “pocket”after the late early Miocene. The strong Mediterra-
nean–Atlantic faunal interchange was cut off in the late Miocene
(Upper Messinian) and started operating again at the base of the Pli-
ocene (Ruggieri, 1967; Hsü et al., 1972). From that time on the Med-
iterranean progressively assumed its present geographical aspect and
underwent a complicated climatic history. Tropical–subtropical con-
ditions were recorded from the central Mediterranean Pliocene, espe-
cially in the lower–early middle part (Raffiet al., 1985; Raffi, 1986;
Monegatti and Raffi, 2001, 2007). In contrast, cooling phases, with a
significant drop in winter temperatures and with summer tempera-
tures closer to the present ones (Raffi, 1986), were reconstructed
for Pleistocene assemblages (particularly from its early part) by
means of the so-called boreal guests (the BGs of authors), namely
North Atlantic molluscs (such as Arctica islandica (Linnaeus), Mya
truncata Linnaeus, Trichotropis borealis G.B. Broderip & Sowerby I,
Buccinum humphreysianum Bennet, Neptunea contraria (Linnaeus)
and so on), which migrated into the Mediterranean through the
Straits of Gibraltar (Gignoux, 1913; Malatesta and Zarlenga, 1986;
Raffi, 1986; Taviani et al., 1991; Di Geronimo et al., 2000). Conditions
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Fig. 2. Warmest (A) and coldest (B) Mediterranean mean monthly sea surface temperatures (°C) in 2008, averaged over August and February, respectively.
From NOAA/ESRL Physical Sciences Division.
100 V. Garilli / Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
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warmer than today are well-known for the late Pleistocene, especially
during MIS 5e, which might be regarded as one of the warmest Qua-
ternary episodes (together with MIS 11, see Zazo et al., 2003a, with
references). At that time, the richest “senegalaise fauna”of Gignoux
(1913), including some tropical West African species such as Polinices
lacteus (Guilding), Strombus latus Gmelin (= S.bubonius Lamarck),
Cantharus viverratus Kiener, Conus ermineus Born (= C.testudinarius
Hwass in Bruguière), Brachidontes puniceus (Gmelin), Cardita senega-
lensis Reeve, etc., migrated into the Mediterranean Sea, colonizing
principally its south-western coasts (Hearty et al., 1986; Zazo et al.,
2003a; Ferranti et al., 2006; Bardají et al., 2009). Despite Ferranti
et al. (2006) confining the presence of S.latus (the most characteristic
species of the last interglacial) to the western Mediterranean MIS 5e,
published data show that this species, or other Senegalese taxa, oc-
curred in the western Mediterranean from MIS 9 or 11 (Balearic Is-
land) and remained afterwards (perhaps up to 60 ka in Italy, see
Zazo et al., 2003a; Bardají et al., 2009, with references). There are
also eastern occurrences of this species in very late Pleistocene
(post-MIS 5e) localities at Nafplion (Zacharias et al., 2000) and
other Tyrrhenian sites from Peloponnesus, the area of Corinth, the
south-central Aegean, including the Aegean arch, and in MIS 5e at Cy-
prus and Israel (Symeonidis and Dermitzakis, 1973; Sivan et al., 1999;
Kéraudren et al., 2000; Theodorou et al., 2005). It is remarkable that
in the Balearic islands the last occurrence of S.latus is at 135 ka, in
MIS 5e, at the end of which a progressive climatic deterioration
started, involving cooler conditions lasting into MIS 5c and 5a.
However, around the Balearics, along the south-western Spanish
coasts, south of the present-day isotherm of 15 °C, S.latus remained
during the entire MIS 5e and later, in MIS 5c/5a. Mean annual SSTs
of around 23–24 °C (no colder than 19–21 °C in winter) and SSS of
34–35 psu have been reconstructed for the western Mediterranean
on the basis of the present-day ecological requirements of the Sene-
galese taxa (Bardají et al., 2009).
3. The tropical eastern Atlantic biogeographical province
This marine bioprovince extends from Cape Blanc (Mauritania) to
Cape Frio (next to the Namibia/Angola border). According to the
hydroclimatic divisions of Le Loeuff and Cosel (1998), this part of
the West African coast is separated into the following regions: north-
ern alternance region (NAR), from Cape Blanc to Cape Verga; western
typical tropical region (WTTR), southward to Cape Palmas; atypical
tropical region (ATR), from Cape Palmas to the Benin front (close to
Cotonou); eastern typical tropical region (ETTR), southward to Cape
Lopez; and southern alternance region (SAR), from Cape Lopez to
Cape Frio (Fig. 4). NAR, ATR and SAR experience periodic upwelling
(Le Loeuff and Cosel, 1998) and their MMSSTs, higher than those in
the Canary Islands, vary respectively from 19–24.5 °C, 26–29 °C, 25–
29 °C and 20.5–26 °C, recorded at their respective geographical bor-
ders (Figs. 4 and 5). WTTR and ETTR have warmer waters and re-
duced SSS (Le Loeuff and Cosel, 1998), with MMSSTs never colder
than about 26 °C (Figs. 4 and 5). In the northernmost part of NAR,
MONTHS
TEMPERATURES °C
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E Med N Creta
E Med N Cyprus
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
SW Mediterranean
W Sicily
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months
sites E Mediterranean
N Creta
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20.5
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Jan
Feb
Mar
Apr
May
Jun
Jul
Aug
Sep
Oct
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Dec
E Mediterranean
N Cyprus
Fig. 3. Mean monthly sea surface temperatures (°C) in selected Mediterranean sites in 2008. Data extrapolated from NOAA/ESRL Physical Sciences Division.
101V. Garilli / Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
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near Cape Blanc, which is the coldest portion of the tropical West
Africa, MMSST is never lower than 20 °C for at least eight months of
the year, whereas southwards to Cape Frio the lowest MMSST of
about 21 °C is recorded for two and four months at Cape Verde and
Cape Frio, respectively (Fig. 5).
In general terms, the modern hydroclimatic setting of the eastern
tropical Atlantic contrasts with that in the Mediterranean and in the
Canary Islands. All the eastern Atlantic tropical regions show a lower
degree of seasonality in comparison with the central-eastern Mediter-
ranean (compare Fig. 3 with Fig. 5). Differences between maximum
and minimum MMSSTs are of about 3–4 °C from Cape Verga to Cape
Lopez. However, some modest analogies can be traced between the
annual variation in MMSSTs in the south-central to eastern Mediterra-
nean and in the northernmost and southernmost tropical regions, and
in particular NAR, where the greatest difference between maximum
and minimum MMSSTs reaches 5.5–8.5 °C (Fig. 5).
Annual salinity averaged over 10–100 m depth generally ranges
from 34 to 35 psu along the entire tropical region, with the exception
of the Gulf of Guinea, in the south-eastern portion of ETTR, where
values of 31–33 psu are recorded. There are no relevant differences
in monthly SSS values (NOAA WOA, 2001).
4. The studied deposits/assemblages
4.1. General notes, recovery information and sampling methods
Shallow-water deposits in Sicily, southern Italy and Greece repre-
sent the main source of investigated molluscan associations. From
this area, the sites of Dattilo, Baglio Inferno–Timpone Pelato, Pizzo
di Core, Cerausi, Grammichele–Catallarga, Cartiera Mulino, Case
Buffa (Sicily) and Kyllini–Kaukalida (NW Peloponnesus, Ionian
Greece) were sampled during several field seasons, in 1998–
2008. For several of them some published data were also considered
(Malatesta, 1960–1963; Curti-Giardina, 1964; Greco, 1970; Ruggieri
and Milone, 1973; Ruggieri and Unti, 1988, with references; Costa,
1989). Other sources, such as for the sites of Tommaso Natale and
Corinth (Aegean Greece), are represented by material housed in the
Ruggieri collection (Museum G.G. Gemmellaro, University of Palermo,
Italy). Further sources of the molluscan associations discussed here
come from the following deposits, for which information was provid-
ed by available literature: Agrigento, Acqua dei Corsari, Belice Valley,
Birgi-Trapani (Sicily), Case Golino, Castrovillari, Crotone, Musalà
(Calabria, southern Italy), Taranto (Puglia, southern Italy), Monastir,
Sfax (Tunisia), Monte Mario, Pisa, Livorno, Santerno (central-north-
ern Italy), Kos and Rhodes (Greece) (Tournoüer, 1876; Fischer,
1877; Cerulli-Irelli, 1907–1916; Gignoux, 1913; Bevilacqua, 1928;
Alberici and Tamini, 1935; Ruggieri, 1949; Nicklès, 1950; Ruggieri,
1953; Malatesta and Nicosia, 1955; Castany et al., 1956; Ruggieri
et al., 1969; Sabelli and Taviani, 1979; Ruggieri et al., 1982 with refer-
ences; Gaetani and Saccà, 1983; Reina, 1985; Ruggieri and Unti, 1988,
with references; Dell'Angelo and Forli, 1995).
As for the sampled deposits, large bulk samples (35–70 l) were
collected, with the exception of the Pizzo di Core, Baglio Inferno–Tim-
pone Pelato and Grammichele–Catallarga sites, of which only very
small, reworked pieces crop out, mainly as a consequence of deep
ploughing. Only handpicking was possible at these sites. Bulk samples
were washed with fresh water in a battery of sieves (mesh 2, 1 and
0.5 mm). Most of the deposits studied during field work are yellowish
to grayish muddy sands containing well-preserved, residual palaeo-
communities (sensu Fagerstrom, 1964). All of the molluscan assem-
blages discussed here belong to the benthic domain from shallow
marine palaeoenvironments. They are considered most useful for cli-
mate reconstruction, as they are highly sensitive to climatic change,
much more so than those from deep, conservative, environments.
The location of the deposits is shown in Fig. 6.
4.2. The Kyllini–Trypiti and Kaukalida successions
By its 500 m-thick Plio-Quaternary evolution, where a wide varie-
ty of palaeoenvironments/assemblages were recorded during more-
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Cape Blanc Cape Blanc
Cape Verde Cape Verde
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Cape Palmas
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Cape Verga
Lobito
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SAR
Frio
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Cape
Frio
SAR
A B
Fig. 4. Coldest mean monthly sea surface temperatures (°C) along West Africa coast in 2008, at Eastern Canary, Cape Blanc and Cape Verde, averaged over January to February (A),
and at Capo Palmas, Cape Lopez and Lobito (B), averaged over August to September. ATR = Atypical tropical region; ETTR = eastern typical tropical region; NAR = northern alter-
nance region; SAR = southern alternance region; WTTR = western typical tropical region.
From NOAA/ESRL Physical Sciences Division.
102 V. Garilli / Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
Author's personal copy
or-less continuous sedimentation, the surroundings of the Kyllini vil-
lage (NW Peloponnesus, Greece, Fig. 7) represent one of the most sat-
isfactory and interesting Mediterranean areas for studying late
Neogene-Quaternary climate change. Notwithstanding this, in-depth
studies of the stratigraphy and palaeontology of this area are lacking.
Four layers bearing thermophilic species have been recognized in the
northern part of the succession, along the cliffed coast between Kyl-
lini and Cape Trypiti: the N2 and H6 layers of Garilli et al. (2005a,
2005b) and Garilli and Galletti (2007), the F5 layer (the P3 of Garilli
et al., 2005b), and the here-named D4 layer (see Fig. 7). The whole
succession is part of an anticline, which originated by a diapiric intru-
sion of Triassic halite and gypsum (Christodoulou, 1971; Hageman,
1976; Kowalczyk and Winter, 1979; Underhill, 1988; Duermeijer et
al., 2000). It consists mainly of sandy-muddy marine-marginal-
lagoon and fresh-water deposits, with a northwards inclination
along the cliff of 40° in the southernmost part and 8–10° in the north-
ern part. The top of the Kyllini succession consists of grayish, muddy,
almost horizontal sands with lagoonal faunal elements and several
remains of earthenware, which are certainly linked to human activi-
ties contemporaneous of the medieval Venetian castle (known as
Paleokastro). Castle ruins are still visible along the coast between Kyl-
lini and Cape Trypiti. This layer is separated by a well-marked ero-
sional surface at the top of yellowish fine sand (see Fig. 7)
containing the gastropod Nassarius musivus (Brocchi), which is repre-
sented by a morph/subspecies bearing delicate sculpture (Nassarius
musivus musivus).
At about 200 m from the Kyllini–Trypiti succession, a short section
cropping out at the small Kaukalida Island (Fig. 7) was also studied in
order to evaluate the possible occurrence of thermophilic molluscan
species. This succession consists mainly of subhorizontal calcarenitic
layers (A–D) with bivalves such as pectinids, glycymerids, abundant
Spondylus gaederopus (Linnaeus) and the coral Cladocora coespitosa
(Linnaeus). This exacoral forms large clumps in layer B. The reddish
layer D also bears gastropods such as Jujubinus striatus (Linnaeus),
J.exasperatus (Pennant), Clanculus corallinus (Gmelin), Columbella
rustica (Linnaeus), and Conus mediterraneus (Hwass in Bruguière).
Layer E consists of a rudite, apparently sterile. Erosional surfaces
occur at the top of layers C and E. A very similar succession, at 1–
4 masl, was observed near the small village of Loutra–Kyllini, about
8 km south of Kyllini.
4.3. The thermophilic species and criteria for recognition
From the Quaternary deposits cited above, 15 taxa have been
identified as thermophilic species that are useful for reconstructing
the climatic and hydrological condition of the south-central and east-
ern parts of the Mediterranean area. A taxonomic list of these species
together with main synonymies/chresonymies, their euro-Mediterra-
nean Tertiary distribution and some remarkable extinction events
according to previous studies is shown in Table 1.
The evaluation of the thermophilic status of the species/assemblage
discussed here has been based on the following criteria: 1) their Mio-
MONTHS
TEMPERATURES °C
Cotonou Cape
Lopez Lobito Cape
Frio
E Canary Cape
Blanc
Cape
Verde
Cape
Verga
Cape
Palmas
19 21 26 28.5 27.5 28 26 23.5
19 21.5 27 29 28 29 27 24.5
20 22 27 29 28.5 29 28 25.5
19.5 22 26 29 29 29 27.5 25.5
20 23 27 29.5 29 28 26.5 24.5
21.5 25.5 28.5 28.5 28 25.5 22.5 21
23 27.5 28 27 26.5 25.5 21.5 20.5
24 26.5 27.5 26.5 26 25 22 20.5
24.5 28.5 28 26.5 26.5 25.5 21.5 20.5
23 28.5 28.5 27 28.5 27 23.5 22
21 26.5 29 28.5 29 27 25.5 24
19.5 23.5 28 29 28.5 27.5 27.5 26
19.5
19
19
19
19.5
21
21.5
21.5
22.5
22
20
20
months
sites
Nouakchott
20
20.5
21
20
20
23
26.5
27.5
28.5
27
24
21
NAR WTTR ATR ETTR SAR
18
19
20
21
22
23
24
25
26
27
28
29
30
31
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Eastern Canary
Cape Blanc
Cape Verte
Cape Verga
Capo Palmas
Cotonou
Cape Lopez
Lobito
Cape Frio
Jan
Feb
Mar
Apr
May
Jun
Jul
Aug
Sep
Oct
Nov
Dec
Nouakchott
Fig. 5. Mean monthly sea surface temperatures (°C) in selected eastern Atlantic sites in 2008 (data extrapolated from NOAA/ESRL Physical Sciences Division) and geographical
boundaries of the West Africa tropical regions of Le Loeuff and Cosel (1998). ATR = Atypical tropical region; ETTR = eastern typical tropical region; NAR = northern alternance
region; SAR = southern alternance region; WTTR = western typical tropical region.
103V. Garilli / Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
Author's personal copy
Pliocene derivation, and therefore their characterization as thermo-
philic “Pliocene resuscitates”(sensu Ruggieri, 1987a, 1987b); 2) their
present-day occurrence in tropical regions; and 3) their close phyloge-
netic linkage to species living intropical regions. The supposed phyloge-
netic linkages are based on morphological resemblances and derive
from the author's personal considerations (such as for the linkage Ana-
dara sp.–Anadara geissei (Dunker in Kobelt)) or from previous studies
(Monegatti and Raffi, 2001; Garilli and Galletti, 2007). Species or their
phylogenetically linked taxa that satisfy the requirements of points 2
and 3 permit an estimation of the SSTs by reference to their present
24
26
4
4,5,6
4
4,6
4,5,6,9
4
8
9
11,12,13,14
11
11
11,12,13,14
12
12
1,17
17
23,29 24
11,12,13
11,12,13
1,7
23,29
23,29
23
23,28?
9 23,29
23,29
24
24
24
24
24
26
3,25,27
26
25
25
25
25
3,25
20
20,21
22
20,21,22
16,17,18
16
15
20,21 10
20
other tropical regions
IO
WA
26
26
26
20 2,19
16
17
18
23-26
20
10
11
12
21
22
8
13-14
27
29
9
15
28
*
*
*
Craspedochiton altavillensis
Rissoina decussata
Niso terebellum
Haedropleura bucciniformis
K. parentalis
Jujubinus ? bullula
Kyllinia marchadi
Terebra corrugata
Anadara geissei
Anadara sp.
Chama crenulata
C. placentina
Plicatula mytilina
Corbula revoluta
MEDITERRANEAN PLEISTOCENE RECENT WEST AFRICA
TROPICAL CLIMATIC REGIONS
LOWER MIDDLE UPPER
NAR WTTR
Taxa
SAR
Ersilia aliceae
ATR
central-east Sicily
west-east Greece south Italy
west Sicily
east Tunisia
south Italy
west Sicily
west Greece
east Tunisia
west Sicily ETTR
west Greece
T. reticularis
Strioterebrum basteroti
S. grayi
central-north Italy south Italy
MIS 5e MIS 5a
W AFRICA
Cape Blanc
Cape Verga
Cape Palmas
Benin Front
Cotonou
Cape
Lopez
Cape
Frio
Cape Verde
30°N
20°N
10°N
0°
10°S
Lobito
RECENT WEST AFRICA
TROPICAL CLIMATIC REGIONS
3-6 7
1
10°E 20° 30°
30°N
40°
NAFRICA
S EUROPE
MEDITERRANEAN
19
2
B
A
C
Fig. 6. Location of the Pleistocene Mediterranean sites studied (A) and the tropical West Africa regions of Le Loeuff and Cosel (1998) and other tropical areas (B); distribution
of selected thermophilic species and close descendants (asterisks) (C). ATR = Atypical tropical region; ETTR = eastern typical tropical region; NAR = northern alternance region;
SAR = southern alternance region; WTTR = western typical tropical region; IO = Indian Ocean; WA = West Atlantic. 1 = Monastir; 2 = Sfax; 3 = Birgi-Trapani; 4 = Dattilo; 5 =
Baglio Inferno–Timpone Pelato; 6 = Pizzo di Core; 7 = Tommaso Natale; 8 = Acqua dei Corsari; 9 = Belice Valley; 10 = Agrigento; 11 = Cerausi; 12 = Grammichele; 13 = Car-
tiera Mulino; 14 = Case Buffa; 15 = Musalà; 16 = Case Golino; 17 = Crotone; 18 = Castrovillari; 19 = Taranto; 20 = Monte Mario; 21 = Tuscany, Pisa and Livorno provinces;
22 = Emilia Romagna (Santerno); 23 = Kyllini layer D4; 24 = Kyllini layer F5; 25 = Kyllini layer H6; 26 = Kyllini layer N2; 27 = Corinth; 28 = Kos Island; 29 = Rhodes Island.
104 V. Garilli / Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
Author's personal copy
Kyllini-Trypiti section (upper part)
HOLOCENE
PLEISTOCENE
Mud
Sand
Kyllini village
Cape Trypiti
Kaukalida Island
Last local occurrence
Rudite
Nannofossil(s) occurrence
Layers bearing thermophlic
mollusks
Fig. 7. Location and simplified evolution of the Kaukalida section and of the northern part of the Kyllini–Trypiti succession (NW Peloponnesus, Greece). Note the stratigraphical
position of the four thermophilic mollusc-bearing layers (D4, F5, H6 and N2) as deduced by nannofossil occurrences (black asterisks) and U/Th dating.
105V. Garilli / Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
Author's personal copy
geographical distributions, and according to actualistic and unifor-
mitarianism approaches. A synoptic overview of the distribution of
the selected species (and related/ancestor taxa) in the Mediterranean
Quaternary, the West Africa tropical regions and other tropical regions
is shown in Fig. 6.
5. Palaeoenvironmental setting
As for the deposits of Birgi-Trapani, Dattilo, Baglio Inferno–
Timpone Pelato, Pizzo di Core, Cerausi, Cartiera Mulino, Case
Buffa and Kyllini (layers N2, H6, F5 and D4), the molluscs are indic-
ative of infralitoral bottoms with the phanerogam Posidonia ocea-
nica (Delile), the HP biocoenosis of Pérès and Picard (1964).In
fact, they are rich in gastropods such as Jujubinus spp., Gibbula
spp., Tricolia spp., Cerithium spp., Rissoidae (mainly Alvania Risso
and Pusillina Monterosato) (see also Curti-Giardina, 1964; Greco,
1970; Ruggieri and Unti, 1988; Costa, 1989), that are herbivo-
rous–detritivorous–depositivorous species characteristic of that
biocoenosis. In most of those deposits, bivalve species such as Lor-
ipes lacteus (Poli), Anadara sp., Plicatula mytilina Philippi and
Chama placentina Defrance were found in life position or with artic-
ulated valves. In some deposits Posidonia leaves and rhizomes are
frequent, and in the Kyllini N2 and H6 layers rhizomes were ob-
served in life position, many of them well-preserved (Figs. 8A–C).
Both these layers, 1.5–2 m thick, represent the evolution of P. ocea-
nica palaeobeds and may be regarded as proxies for investigating
Pleistocene Posidonia palaeo“mattes”. In these deposits, as well
as in those from Dattilo and Cartiera Mulino, the occurrence of
L.lacteus,P.mytilina and abundant Pusillina spp. indicates sheltered
environments with fresh-water input. The Tommaso Natale and
Case Golino deposits formed in a shallow coralligenous environ-
ment (Ruggieri, 1953; Ruggieri and Milone, 1973). The molluscan
association from Corinth includes abundant littoral taxa, especially
trochids, rissoids, carditids, and Spondylus gaederopus (Linnaeus),
suggesting deposition in infralittoral bottoms with vegetation. Cal-
carenitic deposits with pectinids, glycymerids and Cladocora,atthe
Kaukalida site, are interpreted as formed in shallow waters sub-
jected to a relatively strong bottom current. At the same locality
the layer containing gastropods (mainly Jujubinus spp. and Colum-
bella rustica)testifies to a shallow bottom with photophilic
algae and the phanerogam Posidonia (AP-HP biocoenoses of Pérès
and Picard, 1964). Similar conditions are inferred for the deposit
at Crotone as indicated by the fauna described by Ruggieri (1949).
For all of the remaining deposits and their respective associations,
no detailed palaeoecological setting is available, mainly due to the
scarcity of information and/or to the fact that they are the result of
mixed-transported communities (sensu Fagerstrom, 1964). However,
in general terms, on the basis of the published lists of associated taxa,
it is possible that they formed in infra-circalitoral conditions.
6. Stratigraphic setting
6.1. Late Middle–Upper Pleistocene deposits
Three U/Th radiometric dates (Table 2) have been performed
on the coral Cladocora coespitosa from the H6 and N2 layers in the
Kyllini–Trypiti section, and from the biocalcarenitic layer B in the
Kaukalida section. The N2 layer has been dated at 80.5 ±5.2 ka,
allowing correlation with MIS 5a. An age of ≤132.8 ±12 ka, attribut-
ed to layer H6, could be interpreted as affected by post-mortem
Table 1
Taxonomic list of the thermophilic species together with main synonymies/chresonymies, their Euro-Mediterranean Tertiary distribution and some remarkable Euro-Mediterra-
nean extinction events according to previous studies.
Thermophilic species Euro-Mediterranean
Tertiary distribution
Synonymy/chresonomy Remarkable Euro-Mediterranean
extinction events according
to previous studies
References
Polyplacophora
Craspedochiton altavillensis
(Seguenza)
Mio-Pliocene Middle–Upper Pleistocene Garilli et al., 2005b.
Gastropoda
Jujubinus?bullula (Fischer) –Trochus turgidulus
bullula Fischer, Gibbula
bullula (Fischer)
Upper Pleistocene (MIS 5e) Ruggieri, 1949; Ruggieri and Unti, 1988.
Rissoina decussata (Montagu) Mio-Pliocene Rissoina punctostriata
(Talavera)
Bałuk, 2006; Chirli, 2006; Garilli, 2008.
Ersilia aliceae Garilli Pliocene Lower Pleistocene? Garilli, 2004.
Niso terebellum (Dyllwin) Miocene; Pliocene
(Mediterranean)
Niso eburnea Risso Lower Pleistocene (Santernian) Dillwyn, 1817; Raffi, 1986; Chirli, 2009.
Haedropleura bucciniformis
(Bellardi)
Mio-Pliocene Bela bucciniformis
(Bellardi)
Greco, 1970.
Kyllinia parentalis Garilli & Galletti Pliocene Middle–Upper Pleistocene Garilli and Galletti, 2007.
Terebra corrugata Lamarck Mio-Pliocene Terebra acuminata
Borson
Lower Pleistocene (Calabrian) Monegatti and Raffi, 2001, 2007.
T.reticularis (Pecchioli in Sacco) Pliocene Lower Pleistocene (Emilian) Bouchet, 1982; Reina, 1985.
Strioterebrum basteroti (Nyst) Miocene Bouchet, 1981.
Strioterebrum grayi (Smith) Mio-Pliocene Strioterebrum pliocenicum
(Fontannes)
Lower Pleistocene (Emilian) Malatesta, 1974; Ruggieri et al., 1982.
Bivalvia
Anadara sp. –Anadara pectinata
authors, A. pectinata minor
authors, A. breislaky
authors, A. syracusensis
authors
Lower Pleistocene Philippi, 1836; Tournoüer, 1876;
Curti-Giardina, 1964; Greco, 1970;
Costa, 1989.
Chama placentina Defrance Pliocene Lower Pleistocene Gignoux, 1913; Malatesta, 1974;
Ruggieri, 1987b.
Plicatula mytilina Philippi Mio-Pliocene Upper Pleistocene (MIS 5e) Greco, 1986; Ruggieri and Unti, 1988;
Lozouet et al., 2003; Ćorićet al., 2004.
Corbula revoluta (Brocchi) Pliocene Corbula cadenati (Nicklès) Ruggieri, 1987a; Monegatti and Raffi, 2001.
106 V. Garilli / Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
Author's personal copy
alteration, responsible for a strong increase in Th leaching, as indicated
by the low value (0.979 ±0.034) of
234
U/
238
U and, consequently, by the
high ratio
230
Th/
234
U. But, morelikely, the low value of
234
U/
238
Uisat-
tributable to fresh water influences, causing reducing SSS that affected
analyzed coral (see dating discussion in Stamatopoulos et al., 1998).
This agrees with the significant presence of the brackish-water gastro-
pod Pusillina lineolata (Michaud) in layer H6. Furthermore, C. coespitosa
is certainly authoctonous in layer H6, as it was found attached to rhi-
zomes of Posidonia oceanica in life position (Figs. 8C–D). Even in the pre-
sent-dayMediterranean, this exacoral may live in lagoon systemsand in
condition of significant alluvial input (Basset et al., 2006; Peirano et al.
2009 with references), where SSS values can consistently decrease. A
similar condition occurs in the marine marginal-lagoon system in the
bay of Kyllini, where several living colonies of C. coespitosa were ob-
served. On this subject, it is interesting that 3–4 months prolonged cul-
ture in an aquarium showed that this coral tolerates low salinity (down
to 34 psu), although its growth rate was lower than at higher salinity
(Riccardo Rodolfo-Metalpa, personal communication, 2009). Further-
more, the U value (2.910±0.106 ppm) excludes post-mortem Th
leaching (Paola Tuccimei, personal communication, 2004). All these
considerations strongly suggest that the date on the H6 layer is reliable,
and consequently this layer should be correlated with MIS 5e. The de-
posits at Birgi-Trapani, Taranto and Sfax also should be attributed to
MIS 5e (Gignoux, 1913; Nicklès, 1950; Ruggieri and Unti, 1988). The an-
alyzed Cladocora from Kaukalida provided an absolute age of 46± 3ka,
allowing correlation with the very late Pleistocene MIS 3.
With regard to the material from Corinth, it is not possible to spec-
ify a precise stratigraphical setting. However, the lack of extinct
species, with the exception of Jujubinus?bullula, suggests an Upper
Pleistocene Tyrrhenian age.
The deposit at Tommaso Natale (NW Sicily) was attributed to MIS
7 by means of aminostratigraphic analyses (Hearty et al., 1986). Its
molluscan assemblage does not include the Strombus latus, the most
common Senegalese guest occurring in the central part of the Medi-
terranean during MIS 5, and the only reported Senegalese guest is
Cantharus viverratus (Ruggieri and Milone, 1973), whose colonization
of that part of the Mediterranean likely occurred before that of S.latus
and other Senegalese guests. The same stratigraphical setting is likely
for the Tunisian deposit at Monastir, which contains the Senegalese
guest C. viverratus, but with no S. latus (see Ruggieri and Unti, 1988
with references). The same age is attributed to the uppermost terrace
at Crotone (stratotype of the Crotonian stage), following Uranium se-
ries dating and aminostratigraphic data (Mauz and Hassler, 2000,
with references).
6.2. Lower–early Middle Pleistocene deposits
For the stratigraphy of the Lower Pleistocene sites cited hereaf-
ter, the reader is referred to Ruggieri et al. (1984).Aconservative
view of the Pleistocene is here preferred to that ratified by Gibbard
et al. (2010), according to whom the base of this Series/Epoch (the
Pliocene/Pleistocene boundary) coincides with the base of the Gela-
sian stage, at 2.58 Ma.
As indicated by Garilli et al. (2005a, with references), the Kyllini F5
layer should be regarded as late Lower–early Middle Pleistocene in
age, because it underlies by about 40–50 m a turritellid-rich layer
B
C
D
A
Fig. 8. Rhizomes of the phanerogam Posidonia oceanica (Delile). (A, C) From the layer H6 (Kyllini, NW Peloponnesus, late Pleistocene MIS 5e). (B) Recent from Mondello (Palermo,
NW Sicily) for comparison. Note the remains of a Cladocora coespitosa (Linnaeus) colony attached to the rhizome (indicated by white circles). (D) Close up of the dotted circle in C
showing a polyp skeleton of the exacoral preserving internal structure. Scale bars: 1 cm.
Table 2
Results of the U/Th analyses performed on the coral Cladocora coespitosa from the Kyllini–Trypiti and Kaukalida Island successions (NW Peloponnesus, Greece).
Site Th-230/Th-232 U-234/U-238 Th-230/U-234 ppm U Age
(ka)
Kaukalida layer B 68.21 ±15.02 1.120± 0.030 0.030 ±0.020 2.510±0.080 46± 3
Kyllini layer N2 120.95±27.68 1.055±0.032 0.526 ±0.023 2.705±0.093 80.5±5.2
Kyllini layer H6 42±6 0.979±0.034 0.701 ±0.032 2.910±0.106 ≤132 ± 12
107V. Garilli / Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
Author's personal copy
containing the nannofossil association Crenalithus asanoi (Sato &
Takayama) and Gephyrocapsa sp. 3, indicating an age of 0.99–
0.78 Ma. This interpretation appears correct, as the F5 layer overlies
by some tens of meters a muddy layer with C.asanoi, whose Mediter-
ranean first occurrence is recorded at 1.122 Ma (De Kaenel et al.,
1999), in the early Pleistocene Sicilian substage. An upper Emilian
age is suggested for the oldest layer at Kyllini, the D4 layer, due to
its stratigraphical position in the Kyllini-Trypiti succession. In fact,
the D4 layer, underlying the above-mentioned layer with C. asanoi,
overlies by about one meter a lagoonal layer, where the last Kyllini–
Trypiti local occurrence of Micromelania zitteli (Schwartz) is recorded
(see Fig. 7). It is noteworthy that the extinction of this brackish-water
gastropod approximates the top of the continental Villafranchian
stage (Esu et al., 1993), corresponding to the uppermost part of the
Emilian substage.
A Lower Pleistocene, upper Emilian age should be attributed to the
western Sicilian deposits of Baglio Inferno–Timpone Pelato, Dattilo,
Pizzo di Core and Belice Valley and to that at Castrovillari (S Italy)
(Ruggieri et al., 1982; Ruggieri and Unti, 1988). The same age is in-
ferred for the deposit at Cerausi (central Sicily), as its attribution to
the late Pliocene (Greco, 1970) is not correct (Ruggieri, ex littera to
Stefano Palazzi, 1998). A less detailed stratigraphical setting is avail-
able for the deposits of Cartiera Mulino and Case Buffa. A possible
very early Lower Pleistocene (Santernian) age was supposed for the
former deposit by Costa (1989), while an unspecified Lower Pleisto-
cene age was attributed to the latter one (Conti et al., 1979). It cannot
be excluded that both deposits are of Emilian age, as they overlie very
thick calcarenitic deposits containing the boreal guest Arctica islandica
(presumably early Lower Pleistocene). The species reported from
Grammichele–Catallarga are very likely from the same deposit with
terebrids studied by Malatesta (1960–1963). They are from a mixed
palaeocommunity where taxa with different climatic and bathymetric
requirements are found together, due to reworking. Malatesta (1960–
1963) indicated a Calabrian age (Santernian–Emilian substages) for
this deposit. The same age was attributed to the Casa Golino and
Musalà sites, in south Italy (Calabria) (Ruggieri, 1953; Gaetani and
Saccà, 1983), and to the Sicilian Agrigento site (Malatesta and Nicosia,
1955). A Lower Pleistocene age should also be attributed to the Kos
and Rhodes deposits as indicated by their respective molluscan as-
semblages (Tournoüer, 1876; Fischer, 1877).
The reported specimens of Terebra reticularis were handpicked by
Reina (1985) from dug muddy sediments at Cava Fazio, also known as
Cava Puleo (Acqua dei Corsari, Ficarazzi, Palermo, northwestern Sici-
ly). Consequently, the stratigraphical context of this finding is lost.
Very likely these specimens are not from the well-known bluish
clays with boreal guests of Sicilian age cropping out at this locality.
As Reina himself indicated, they could be considered as coming
from older, Emilian sediments. This is probable, as late Emilian sedi-
ments effectively underlie the Sicilian clays in boreholes at this local-
ity, or in the immediate vicinity (Di Stefano and Rio, 1980).
7. The thermophilic species: results
7.1. The index species
By means of their Mediterranean Pleistocene distribution, among
the thermophilic taxa listed in Table 1 and Fig. 6,Anadara sp. and Juju-
binus?bullula (Fig. 9) appear to be the most common species in the
Mediterranean Pleistocene sites discussed here. They therefore repre-
sent an index taxonomic component of the specified interglacial epi-
sodes, above all in reconstructed conditions that can be related to
Posidonia oceanica palaeoenvironments. Notwithstanding this, these
species are not well-known in the palaeontological literature. In par-
ticular Anadara sp. has often been misunderstood (see chresonomy in
Table 1). J.? bullula could be considered a Mediterranean cryptogenic
species, although there are very few similar upper Tertiary species,
namely Trochus turgidulus Brocchi and Gibbula simulans (De Stefani
& Pantanelli). The derivation of Anadara sp. might be traced to some
Mio-Pliocene Euro–Mediterranean taxa, such as Arca pectinata Broc-
chi and A.burdigalina Mayer. However, Anadara sp. appears phyloge-
netically linked to the West African living species Anadara geissei
(Dunker in Kobelt), from which only few taxonomic differences can
be noted (compare Fig. 9A with Fig. 9C).
7.2. The palaeoclimatic significance
The thermophilic species listed in Table 1 and Fig. 6 can be
grouped into 4 stocks according to their palaeo- and modern geo-
graphical ranges and their use as proxy data for recognizing Mediter-
ranean Pleistocene interglacials and evaluating palaeo-SSTs.
1) A stock including Craspedochiton altavillensis and Haedropleura
bucciniformis, namely Euro-Mediterranean (and Paratethyan) ex-
tinct species, whose Mio-Pliocene common occurrence (Greco, 1970;
Garilli et al., 2005b) could be regarded as the main indicator of their
thermophilic significance (“Pliocene resuscitates”of Ruggieri, 1987a,
1987b). They are useful for recognizing warm climatic events, but
they are not useful for the recognition of palaeo-SSTs. 2) A second
stock with Ersilia aliceae,Strioterebrum basteroti and Plicatula mytilina,
namely species that are indicative of warm climatic conditions based
on the distribution of their respective genus/family. Also these species
occurred in the Mio-Pliocene of the Euro-Mediterranean region, a pe-
riod that is commonly considered as characterized by warmer climatic
conditions than the present-day Mediterranean (Bouchet, 1981;
Greco, 1986; Lozouet et al., 2003; Ćorićet al., 2004; Garilli, 2004). Pli-
catula mytilina belongs to a strictly tropical shallow-water genus, ab-
sent from the present-day Mediterranean (Cox and Hertlein, 1969;
Squires and Saul, 1997; Harzhauser et al., 2003) and represented
today by few species. For example, Plicatula australis Lamarck and
P.tuberculosa Nomura live in Malaysia-W Australia (data from the
Western Australian Museum, 2006) and China and Taiwan (Bernard
et al., 1993), respectively, whereas the few North American extant
AB
C
E
D
Fig. 9. The most representative species of the thermophilic associations studied here,
Jujubinus?bullula (Fischer) and Anadara sp., and Anadara geissei (Dunker in Kobelt),
considered phylogenetically close to the latter species. (A–B) Anadara sp. showing ex-
terior of the left valve (A) and dorsal view (B) of the same specimen (29 mm in width)
from the late Pleistocene MIS 5a N2 layer, Kyllini, NW Peloponnesus, Greece. (C) Ana-
dara geissei, exterior of the left valve (71.1 mm in width), recent from Pointe Noire,
French Congo (© 2011 Guido T. and Philippe Poppe —www.conchology.be). (D) Syn-
type of Trochus bullula Fischer (12 mm in height), number R07497, Muséum National
d'Histoire Naturelle, Département Histoire de la Terre, Paris, d'Orbigny collection, Pleis-
tocene of Rhodes, Greece. (E) Jujubinus?bullula (14.1 mm in height) from the N2 layer,
Kyllini, NW Peloponnesus, Greece.
108 V. Garilli / Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
Author's personal copy
species are from the southern Gulf of California, two of them ranging
to Ecuador (Keen, 1971), and from the Gulf of Mexico, including Flor-
ida, to Bermuda (Database of Western Atlantic Marine Mollusca). The
only species from West Africa, P.angolensis Cosel, lives from Cameroon
to northern Angola, at Luanda (Cosel, 1995). As a whole, MMSSTs of
these regions are never colder than about 19.5–20 °C (NOAA-ESRL,cal-
culated over the year 2008). E. aliceae belongstoagenusthatisconsid-
ered to be a tropical–subtropical taxon, represented today by E.stancyki
Warén, from Florida (Warén, 1980, 1984), and E.mediterranea (Monter-
osato), which is consistently limited to the south-central and eastern-
most parts of the present-day Mediterranean (Ghisotti, 1978; Garilli,
2004). For the palaeoclimatic significance of the terebrid S. basteroti
see below. 3) A stock with Kyllinia parentalis,Anadara sp. and Chama
placentina, whose thermophilic significance comes from the distribu-
tion of their respective closely related modern species, namely Kyllinia
marchadi,Anadara geissei and Chama crenulata.Kyllinia marchadi (the
only known living representative of the genus Kyllinia) lives in the
southern part of WTTR and in the central part of SAR (Rolán et al.,
1998,asDiaugasma marchadi;Garilli and Galletti, 2007)inclimaticcon-
ditions with lowest MMSSTs of 21.5 °C and MMSSTs warmer than 24 °C
for at least 7 months of the year. Anadara geissei is known from all the
West African tropical regions with the exception of ETTR (Nicklès,
1950; Collignon, 1960; Le Loeuff, 1993; Ardovini and Cossignani,
2004). It therefore tolerates minimum temperatures not colder than
19 °C and requires MMSSTs ≥20 °C for at least 8 months of the year.
The same palaeoclimatic significance can be inferred from the distribu-
tion of C.crenulata, which lives in NAR, ETTR and SAR (Nicklès, 1950;
Monegatti and Raffi,2001). 4) The fourth stock includes Rissoina decus-
sata,Niso terebellum,Strioterebrum grayi,Terebra corrugata,Terebra reti-
cularis and Corbula revoluta, namely species living today in fully tropical
conditions. R.decussata occurs in all the West African tropical regions,
but it is not present north of Nouakchott (18°04′N, 16°W) (Gofas,
1999,asRissoina punctostriata). It tolerates minimum MMSSTs of
20 °C and requires MMSSTs warmer than 20 °C for at least 9 months
of the year. As a whole, similar climatic conditions are also recorded in
the Caribbean area and in the Gulf of Mexico where this species also
lives (Arnow et al., 1963; Ponder, 1985; Prieto et al., 2003; Narciso et
al., 2005; Database of Western Atlantic Marine Mollusca). According
to Davoli (1976), the Atlantic terebrids (a typical tropical group) do
not tolerate SSTs lower than 18 °C and need SSTs higher than 20 °C for
6–9 months of the year. This hydroclimatic setting effectively matches
the requirements of S. grayi,T. corrugata and T.reticularis, which are
overall distributed south of Cape Blanc to the Angolan coasts, from
NAR to SAR regions (Bouchet, 1982; Ardovini and Cossignani, 2004).
Niso terebellum belongs to a genus parasitic on Echinoidermata, which
today is restricted to the temperate–tropical provinces (Bouchet and
Warén, 1986). Most Niso species live in the Indo-Pacific, the Caribbean
area and the Gulf of Mexico (Bouchet and Warén, 1986; Appeltans et
al., 2011). Along the West African coast it is represented by N.chevreuxi
Dautzenberg, living in Mauritania, Cape Verde archipelago, Senegal,
Gabon and Angola (Appeltans et al., 2011). At the Nicobar Islands,
type locality of N.terebellum (see Dillwyn, 1817, p. 873), climatic condi-
tions are characterized by a very low seasonality, with MMSSTs of 28.5–
29.5 °C (NOAA-ESRL). These climatic conditions do not match exactly
the effective temperature requirements of this species, which seems,
as well as other parasitic molluscs, to be euribathyal according to tem-
perature gradients and/or bathymetric requirements of its host. Corbula
revoluta have a very similar palaeoclimatic significance to that shown
by R.decussata but, since it does not occur north of Cape Verde
(14°45′N17°25
′W), having being recorded from Cape Verde and Cape
Palmas and on the coasts of Cameroon (Giresse et al., 1996,asCorbula
cadenati (Nicklès); Monegatti and Raffi,2001), tolerates a minimum
MMSST of 21 °C.
A different role is played by Jujubinus?bullula, the climatic signif-
icance of which derives from the fact that it always occurs with ther-
mophilic species/associations.
8. Discussion
8.1. Palaeoecology, palaeohydrological, and salinity conditions
The results of this study indicate that there is a strong linkage be-
tween biodiversity changes within marine shallow-watermolluscan as-
semblages from the south-central to eastern Mediterranean and the
onset of warmer climatic conditions throughout the Lower–Middle
and late Pleistocene (MIS 5e and 5a) history of this basin. As for the
late Pleistocene interglacial MIS 5, the discussed assemblages represent
thermophilic stocks that are absolutely original in comparison to those
known as the Senegalese fauna. As for the Lower–Middle Pleistocene,
the studied species are the most exhaustive molluscan assemblages
allowing estimation of SSTs during interglacials, with special regard to
that which occurred at the top of the Emilian substage, namely the
“Emilian transgression”of Ruggieri (1978, 1987b). This event is corre-
lated with the MIS 37, which according to δ
18
O stratigraphy (Gibbard
and Kolfschoten van, 2004) appears to be the warmest peak in the up-
permost part of the Emilian substage, at about 1.25 Ma (Sprovieri,
1993).
As a whole, the assemblages discussed here permit outlining cli-
matic conditions characterized by SSTs never lower than 19–21 °C,
with values ≥20 °C for at least 8 months of the year and ≥24° for at
least 5–6 months. Comparable conditions regarding February SSTs
were reported for the mid-Pliocene of western Iberia, at the latitude
of Mondego (Silva et al., 2010), in contrast with the PRISM2 Project
data set (Dowsett et al., 1999), which indicates 17–18 °C for the
same locality. Assuming that the highest values of SSTs in the past
Mediterranean did not differ from those of today, the studied assem-
blages therefore suggest that less marked seasonality regimes were
manifested during the warm Mediterranean Pleistocene episodes.
This contrasts with the climatic settings delineated for Pleistocene
cold events in the same basin, where SSTs were lower than 9–10 °C
in February and not lower than 19–20 °C for at least 3–4 months
(Raffi, 1986). In particular, all the selected species living today along
the coasts of West Africa, and their phyletically close species, with
the exception of Chama crenulata, do not occur north of Cape Verde-
Nouakchott, indicating the requirement of a climatic regime with
SSTs ≥24 °C for at least 5–6 months of the year. This situation is com-
parable only with the easternmost part of the Mediterranean, where
today most of the tropical Lessepsian molluscs have well-established
populations.
The highest concentration of the selected species is from the
Lower Pleistocene of Sicily, where 13 out of the 15 selected species
occur. Only 6 and 4 thermophilic taxa occur in Lower Pleistocene de-
posits of central and northern Italy, respectively. Nine species overall
are from the Lower–early Middle Pleistocene of Greece (Kyllini
and Rhodes) and from interglacial MIS 5, in the upper part of the
Kyllini–Trypiti succession. All this allows stressing that warm con-
dition recorded here from the Lower Pleistocene should be consid-
ered to be of the same climatic order as (possibly slightly warmer
than?) those that occurred during the MIS 5e. However, the numer-
ous glacial events that occurred throughout the Quaternary very
likely played a role in triggering the apparent decreasing biodiversi-
ty trend that affected the thermophilic associations during the early
to late Pleistocene. Six thermophilic species are recorded from the
Greek MIS 5e and MIS 5a (H6 and N2 layers at Kyllini, respectively).
Even though no terebrid species occurs in the latter stage, the pres-
ence of unquestionably warm species, such as Chama placentina and
Plicatula mytilina, indicates that warm condition clearly were main-
tained in eastern Mediterranean during MIS 5a, at 80.5 ka. This con-
trasts with the northern hemisphere and the north-western
Mediterranean climate data sets indicating a progressive deteriora-
tion following the warmest peak of MIS 5e, and involving MIS 5a
(Bardají et al., 2009). As for the late Middle Pleistocene central
Mediterranean MIS 7, only 3 thermophilic species have been
109V. Garilli / Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
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found. However, all the middle-late portion of the Middle Pleisto-
cene (MIS 11 to MIS 7) is scarcely known in the central to eastern
Mediterranean, at least with regard to information on the mollus-
can fauna. Concerning this, there are some promising deposits,
dated MIS 11 and MIS 7, in the northernmost part of the western
Peloponnesus (Stamatopoulos et al., 1998), whose molluscan
faunas are worth studying deeply. Also, the Pleistocene molluscs
from the easternmost Mediterranean lack an in-depth study that
could greatly increase the number of thermophilic species from
that part of the basin.
Selected thermophilic species, or their close descendants, living
today along the tropical West African coast, require a SSS value not
over 34–35 psu, which is consistently lower than the average value
(38 psu) in the Mediterranean, where 36 and 40 psu are recorded in
the westernmost and easternmost part, respectively. This suggests
that during the warm episodes analyzed from the south-central to
eastern Mediterranean, the SSS was consistently lower than today,
possibly due to an increase in rainfall/runoff. This matches with re-
sults obtained for MIS 5 from the western Mediterranean, in the
Alboran Sea and along the south-central Spanish coasts, where SSSs
of 34–35 psu were reconstructed by means of the Senegalese fauna
(Pérez Folgado et al., 2004; Bardají et al., 2009). Reducing salinity
during the warm events investigated here might find support from
the low
234
U/
238
U values (0.979 and 1.055 in the H6 and N2 layers, re-
spectively), which in some way is linked to the salinity values of the
water from which the carbonate is assimilated by skeletal organisms
(Paola Tuccimei personal communication, 2004). This ratio is consis-
tently higher (1.120) in the Cladocora specimens dated from the Kau-
kalida section (MIS 3), where molluscan assemblages indicate climate
conditions comparable to those in the present-day Mediterranean.
However, different variables should be evaluated to confirm this
assumption.
Most of the selected species are linked to shallow palaeobottoms
with Posidonia oceanica. This relationship is so strong for Jujubinus?
bullula and Anadara sp., these taxa having been recorded almost ex-
clusively from that palaeoenvironment, that these taxa should be
considered preferential (sensu Pérès and Picard, 1964) of Pleistocene
Mediterranean Posidonia beds. The occurrence of P. oceanica in
palaeoenvironments with interpreted salinity near to 34–35 psu, con-
siderably lower than that in places where the seagrass is endemic at
present, should not be viewed with suspicion. In fact P. oceanica toler-
ates hyposaline waters of 25–36.4 psu (Fernández-Torquemada and
Sánchez-Lizaso, 2005), but undergoes stress from hypersalinity with
values greater than 38–38.5 psu (approximately mean SSS in Medi-
terranean) (Latorre, 2005; Sánchez-Lizaso et al., 2008; Ruíz et al.,
2009).
8.2. Climate interpretation of the warmer past: a predictive role?
Climatologists agree in regarding global warming as the naked
truth. They also consider the present-day Mediterranean as one of
the most sensitive areas for prediction of future climate conditions.
It is therefore obvious that expectations from the climate reconstruc-
tions of the warmer Mediterranean past, as a representative interval
of natural climate variability, are high. However, while the Mediterra-
nean warm palaeoclimate settings interpreted here indicate the oc-
currence of hydrological conditions characterized by an increase in
rainfall/runoff, climatic projections outlined as a consequence of the
global warming indicate a decrease in precipitation in south Europe
and the Mediterranean, with particular regard to the summer months
in southern and eastern regions (Oikonomou et al., 2008; Plan Bleu,
2010). Probably this discrepancy between past reconstructions and
future projections is due to the role played by human activities
(greenhouse gases emission) in influencing and/or accelerating cli-
mate change. Be that as it may, the climatic reconstructions of warm-
er recent past add important information to our understanding and
consciousness of how human activity can shift the course of natural
climate change.
8.3. Palaeobiogeography
The association Jujubinus?bullula–Anadara sp., the most represen-
tatives species within the studied assemblages, appears limited to the
south-central to eastern Mediterranean basin, never having being
recorded north of about 40° N (northern Calabria, southern Italy)
and west of 10°50′E (eastern Tunisia). Anadara sp. shows a more re-
stricted distribution between about 38° N (western Sicily and north-
western Peloponnesus) and about 12°40′E (western Sicily) (see
Fig. 10); the same distribution is shown even by the four species of
the strictly tropical family Terebridae found in the Mediterranean
Pleistocene. All this suggests that the south-central to eastern Medi-
terranean was, as at the present time, a palaeogeographical region
warmer than the western part of the basin, and indicates that the
boundary between the two (palaeo)Mediterranean subregions
matches with that reported by Bianchi (2007, with references) for
the present-day Sicilian–Tunisian Strait (red curve a in Fig. 10).
As indicated by Zazo et al. (2003b), the Cape Verde Archipelago
should be regarded as the main source of “Senegalese”taxa, whose
oldest occurrence is from a Cape Verde terrace, dated at 800 ka (the
oldest age of the Senegalese assemblage from Atlantic deposits,
according to the available data). This indicates that the direction of
migration of the Senegalese assemblages towards the Mediterranean,
occurring during the Middle to Upper Pleistocene interglacials
(mainly MIS 5e), was very likely persistently unidirectional. A differ-
ent regime of migration, with an opposite direction (from the Medi-
terranean towards the Atlantic) probably occurred for several Mio-
Pliocene taxa as supported by what is indicated by Le Loeuff and
Cosel (1998), who recognized species from the Italian Pliocene, or
their close descendants, in some refuge areas (mainly Southern An-
gola, and the coast between Cape Verde and part of the continental
shelf off Casamance) where the MMSSTs are never cooler than
24 °C for at least six months of the year (Monegatti and Raffi,
2001). The results of this study indicate that the source for the Med-
iterranean Quaternary thermophilic associations did not necessary
derive from the eastern Atlantic tropical regions as appears from
the actual evidence (the Senegalese guests). Taxa such as the
(Euro)Mediterranean Upper Cainozoic–Upper Pleistocene species
Kyllinia parentalis,Rissoina decussata,Terebra corrugata,Terebra reti-
cularis,Strioterebrum grayi,Corbula revoluta and Chama placentina,
living in the tropical African regions, or represented there by close
descendants, seem to have originated from Mediterranean area. Ef-
fectively, no fossil Atlantic record, at least older than those in Medi-
terranean, is available for these species. Even a strictly endemic
assemblage from the Mediterranean played a considerable role in
characterizing warm Quaternary Mediterranean episodes: Craspedo-
chiton altavillensis,Ersilia aliceae,Strioterebrum basteroti,Plicatula
mytilina, and above all Jujubinus?bullula and Anadara sp.
8.4. Stratigraphy of the Kyllini succession and disappearance events
within selected species
The U/Th dating presented here rewrites the stratigraphy of the NE
part of the Peloponnesus, with particular regard to the northern part
of the Kylliny–Trypiti succession and to the Kaukalida section, which
previously were attributed to the Plio-early Pleistocene (Christodoulou,
1971,Pliocene;Hageman, 1976,earlyPleistocene;Kowalczyk
and Winter, 1979, late Pliocene–early Pleistocene; Duermeijer et al.,
2000, Pliocene) and to the late Pleistocene, Tyrrhenian (Kowalczyk
and Winter, 1979), respectively. Whereas the chronology of the
Kyllini–Trypiti succession should be extended at least to MIS 5a,
the age of the subhorizontal deposits at Kaukalida (together with
similar neighboring coastal deposits) is younger and is attributed
110 V. Garilli / Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
Author's personal copy
to MIS 3. The youngest substage of the MIS 5 is, therefore, recorded
for the first time in the Peloponnesus, where only two interstadials,
MIS 5e and MIS 5c, were previously reported (Kéraudren et al.,
2000).
This data set also allows accepting a later date for the extinction of
taxa previously thought to have become extinct in the (Mediterranean)
Lower–Middle Pleistocene. The Mediterranean extinction of the family
Terebridae is to be extended to MIS 5e, where Terebra corrugata has
been found in layer H6 at Kyllini. The same data set also allows extend-
ing the last appearance of Craspedochiton altavillensis,Jujubinus?bullula,
Ersilia aliceae (MIS 5a), Niso terebellum (Mediterranean extinction in the
Emilian substage), Anadara sp., Chama placentina,Plicatula mytilina
(MIS 5a) and Kyllinia parentalis (MIS 5e). Furthermore, the warm-tem-
perate Mediterranean Mio-Pliocene gastropod (Malatesta, 1974)Nas-
sarius musivus musivus, previously used as a marker for distinguishing
Pliocene from Lower Pleistocene (Buccheri, 1970), should be regarded
with suspicion, having being recorded in Greek deposits younger than
MIS 5a (Kyllini–Trypiti succession) and older than the MIS 3 (Kauka-
lida). Therefore, although several glacial Pleistocene events influenced
the history of thermophilic molluscan assemblages, causing the (local)
disappearance of several species, many warm selected taxa survived
in the eastern Mediterranean up to MIS 5a.
9. Conclusion
Fifteen shallow-water species, representing thermophilic molluscan
associations absolutely original in comparison to those previously
known as the Senegalese fauna, have been successfully used for palaeo-
climatic reconstruction of some Pleistocene warm episodes in the
south-central and eastern Mediterranean. By means of their palaeocli-
matic significance, climatic conditions have been interpreted with win-
ter MMSSTs 4–6 °C and at least 2–4 °C warmer than in the present-day
western Sicily and easternmost part of the Mediterranean Sea, respec-
tively. Climatic regime with SSTs ≥24 °C for at least 5–6months of
the year and ≥20 °C for at least 8 months of the year wasalso indicated.
These conditions have been recognized in the Lower Pleistocene, with
particular regard to the Emilian substage, in the Lower–early Middle
Pleistocene and in the Upper Pleistocene MIS 5e and MIS 5a. Similar
conditions are inferred to MIS 7 from Sicily and southern Italy, in spite
of fewer thermophilic species having been detected there (possibly
due to the scarcity of investigated deposits and partly to the lack of in-
depth studies of their molluscan assemblages). In contrast with glacial
Pleistocene events, the detected warm episodes are characterized by a
lower degree of seasonality, with a difference between summer and
winter SSTs of 6–7°C(vs.10–11 °C recorded in the present-day central
to eastern Mediterranean).
No remarkable climatic differences have been observed between
MIS 5e (132.8 ka) and MIS 5a (80.5 ka) as recognized in the north-
western Peloponnesus (Kyllini) by U/Th radiometric analysis per-
formed of Cladocora coespitosa. This suggests that, at least in those
areas, climatic conditions appreciably warmer than today persisted
at the end of MIS 5. No warm species has been found in the 46 ka
layer at Kaukalida, deposited during the very late Pleistocene MIS 3.
The U/Th dating outlines a new stratigraphical setting for the
Kyllini–Trypiti succession (and the NW Peloponnesus): the north-
ern part of this succession is now attributed at least to a Lower
Pleistocene–Upper Pleistocene (MIS 5a) range.
Molluscan assemblages discussed here indicate that during
detected warm Pleistocene episodes, south-central to eastern Medi-
terranean SSS was consistently lower than today, with values close
to those now recorded in tropical West Africa (34–35 psu vs. 37–
38 psu and 39–40 psu in the south-central and eastern Mediterra-
nean, respectively). A different Mediterranean hydrological regime,
triggered by increased in rainfall/runoff, therefore was manifested
during warmer Pleistocene periods.
Data from selected and discussed associations show that mollus-
can biodiversity responses in central to eastern Mediterranean warm-
er Quaternary episodes were not always linked to the migration of
taxa from West Africa, but that an endemic component from the
Euro-Mediterranean realm played an important role in characterizing
these episodes.
The distribution of some of the most meaningful studied species
indicates that, during the Pleistocene as it is today, the Mediterranean
Sea was divided into western cooler and eastern warmer palaeobio-
provinces, with a boundary zone possibly located in the proximities
of the current Sicilian–Tunisian Strait.
Fig. 10. Pleistocene Mediterranean distribution of the thermophilic taxa Jujubinus?bullula,Anadara sp. and Terebridae as deduced from literature and data from the present article
(see legend in figure). The red curves indicate the position of the present-day western-eastern Mediterranean boundary according to different authors (from Bianchi, 2007, with
references). The blue dotted line delimits the distribution of J.?bullula–Anadara sp. and that of Pleistocene Mediterranean Terebridae, suggesting the hypothetical border of the
hydroclimatic eastern Mediterranean region during the Pleistocene. 1 = possible late Middle Pleistocene of Monastir; 2 = Upper Pleistocene Tyrrhenian of Birgi-Trapani; 3 = Emi-
lian of Dattilo, Pizzo di Core e Timpone Pelato; 4 = Emilian of Ficarazzi; 5 = Emilian of the Belice Valley 6 = Emilian of Cerausi 7 = Possible Emilian of Grammichele; 8 = Possible
Emilian of Cartiera Mulino and Case Buffa; 9 = Lower Pleistocene of Musalà; 10 = Lower Pleistocene of Case Golino; 11 = late Middle Pleistocene of Crotone; 12 = Emilian of
Castrovillari; 13 = Lower to Upper Pleistocene of Kyllini; 14 = late Pleistocene of the Corinth area; 15 = probable Lower Pleistocene of Kos; 16 = Lower Pleistocene of Rhodes.
111V. Garilli / Palaeogeography, Palaeoclimatology, Palaeoecology 312 (2011) 98–114
Author's personal copy
Acknowledgments
This paper would not have been possible without the generous ef-
fort of friends and colleagues. Eugenio Di Liberto (Bagheria, Italy),
Giacomo Gullo (Partinico, Italy), Luca Galletti (APEMA, Palermo, Italy),
Stefano Palazzi (Milo, Italy), Francesco Pollina (APEMA) and Agatino
Reitano (Tremestieri, Italy) provided assistance during several field sea-
sons. L. Galletti also provided images in Figs. 8A–B. Guido T. and Philippe
Poppe (Conchology, Inc., Cebu Light Industrial Park, Mactan Island,
Philippines) kindly provided permission to republish images in
Fig. 9C. Luca Bertolaso (Correggio, Italy), L. Galletti, S. Palazzi, Carlos
Marques da Silva (Centro e Departamento de Geologia da Faculdade
de Ciências, University of Lisboa, Portugal), Barbara Mauz (Department
of Geography, University of Liverpool, UK), Riccardo Rodolfo-Metalpa
(Centre Scientifique de Monaco, Principality of Monaco), Paola Tuccimei
(Università Roma Tre, Roma, Italy), and Evi Vardala-Theodorou
(Goulandris Natural History Museum, Kifissia, Greece) are thanked for
the helpfuldiscussionson various itemsand/or for providing literature.
S. Palazzi also made available his correspondence with Giuliano
Ruggieri. Field workings at Kyllini in 2000 and 2002–2004 were partial-
ly funded by MURST 60% and by European Community (contract n.
ENV4-CT98-5125). U/Th dating was possible thanks to MURST doctoral
funds to the writer. NOAA_OI_SST_V2 data were provided by the
NOAA/OAR/ESRL PSD, Boulder, Colorado, USA, from http://www.esrl.
noaa.gov/psd/. Didier Merle and Jean-Michel Pacaud (Muséum National
d'Histoire Naturelle, Paris, France) allowed me to visit the collection
d'Orbigny housing the syntype of Trochus bullula. Thanks are also due
to Marina L. Aguirre (CONICET, INGEA UNLP, La Plata, Argentina) and
Alan G. Beu (GNS Scienze, Lower Hutt, New Zealand) for their construc-
tive comments.
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