Geobios 40 (2007) 325–337

http://france.elsevier.com/direct/GEOBIO

Original article
Paratethyan Ostracod immigrants in Italy during the Late Miocene
Immigrants paratéthysiens parmi les ostracodes au
Miocène supérieur en Italie
Elsa Gliozzi a,b,*, Maria Elena Ceci a, Francesco Grossi a, Silvia Ligios a
a
Dipartimento di Scienze Geologiche, Università Roma Tre, Largo S. Leonardo Murialdo, 1, 00145 Roma, Italy
b
IGAG, CNR, c/o Dipartimento di Scienze della Terra, Università La Sapienza, Roma, Italy
Received 18 January 2005; accepted 27 October 2006
Available online 1 May 2007

Abstract
In this paper the ostracod assemblages recovered from several brackish Late Miocene Italian deposits have been analysed from a
palaeobiogeographical perspective. During late Tortonian-early Messinian it is possible to recognize in Italy rich ostracod assemblages
characterized by a wide contingent of taxa with central European or Mediterranean affinity, while only few brackish and freshwater ostracods
show Paratethyan affinity. The recognized composition of the ostracod assemblages matches the palaeogeographic setting of the palaeo-
Mediterranean/Paratethys at that moment. In fact during late Tortonian-early Messinian the palaeo-Mediterranean and Paratethysian domains were
divided and, even if the connection via the present Marmara Sea-Strimon Basin was still open, the different salinity between them represented an
ecological barrier, preventing faunal exchanges. Since normal aquatic migration was impossible, it must be assumed that the Paratethyan-like taxa
entered the palaeo-Mediterranean area via passive dispersal by aquatic birds. On the contrary, the ostracod assemblages from the Italian Lago-Mare
deposits show the absolute predominance of Paratethyan taxa, which, according to the known palaeogeographic setting during the late Messinian
Lago-Mare event, could actively migrate from the Paratethys domain, colonizing the palaeo-Mediterranean, whose endemic fauna was severely
impoverished by the Messinian salinity crisis and the following water dilution.
# 2007 Elsevier Masson SAS. All rights reserved.

Résumé
Cette étude est focalisée sur les implications paléobiogéographiques des associations d’ostracodes récoltées sur de nombreux dépôts saumâtres
du Miocène supérieur d’Italie. Au Tortonien supérieur-Messinien inférieur il est possible de reconnaı̂tre en Italie beaucoup d’ostracodes
caractérisés par une affinité méditerranéenne et européenne centrale. Seules quelques ostracodes saumâtres montrent une affinité paratéthysienne.
L’association d’ostracodes retrouvés reflète la géographie de la paléo-Méditerranée et de la Paratéthys à ce moment. Pendant le Tortonien
supérieur-Messinien inférieur, la connexion à travers l’actuelle mer de Marmara et le bassin du Strymon était encore ouverte, mais la différence de
salinité entre la paléo-Méditerranée et la Paratéthys représentait une barrière écologique interdisant les échanges faunistiques à l’origine des deux
bioprovinces paléo-méditerranéenne à l’Ouest et paratéthysienne à l’Est. Dans ce cas, seule une dispersion passive par les oiseaux aquatiques peut
être envisagée. Au contraire, les ostracodes des dépôts du Lago-Mare d’Italie indiquent la dominance absolue des taxa paratéthysiens : la barrière
écologique entre la paléo-Méditerranée et la Paratéthys avait disparu et cet important contingent d’ostracodes migra effectivement vers l’Ouest, en
colonisant la paléo-Méditerranée dont la faune endémique avait été fortement appauvrie par la crise de salinité messinienne et la dilution des ses
eaux qui s’en suivit.
# 2007 Elsevier Masson SAS. All rights reserved.
Keywords: Late Miocene; Palaeo-Mediterranean; Paratethys; Brackish ostracods; Palaeobiogeography; Dispersal modes

Mots clés : Miocène supérieur ; Paléo-Méditerranée ; Paratéthys ; Ostracodes saumâtres ; Paléobiogéographie ; Modalités de dispersion

1. Introduction

* Corresponding author. During the Miocene the Mediterranean area was affected by
E-mail address: gliozzi@uniroma3.it (E. Gliozzi). important palaeogeographic changes linked to the collision

0016-6995/$ – see front matter # 2007 Elsevier Masson SAS. All rights reserved.
doi:10.1016/j.geobios.2006.10.004
326 E. Gliozzi et al. / Geobios 40 (2007) 325–337

between the African and Adria plates and the subsequent
building of the peri-Mediterranean orogenic belt. During the
Serravallian the emergence of the Carpathian and Alpine
foreland domains progressively closed the northern connection
between the Western (palaeo-Mediterranean) and the Eastern
Tethys (Paratethys), which, from the Burdigalian, was open
through the Pannonian back-arc basin (Meulenkamp and
Sissingh, 2003, and references therein). From this moment, the
Paratethyan marine realm was separated into several brackish
(oligo-mesohaline) basins (Pannonian, with its Vienna and
Styrian sub-basins, Dacic, Euxinic and Caspian-Aral basins),
which, notwithstanding the subsequent uplift of the Dinarid and
Balkan peri-Mediterranean chains, maintained a partial/
intermittent connection among them. Within these basins,
molluscs and ostracods underwent an important adaptive
radiation, giving rise to the Paratethyan palaeobioprovince,
distinctly different from the palaeo-Mediterranean palaeobio-
province (Marinescu, 1992) (Fig. 1).
During the Tortonian and early Messinian the palaeo-
Mediterranean was connected with the Atlantic Ocean, and,
consequently, it was characterized by fully marine conditions.
Although not all authors agree (Orszag-Sperber et al., 2000),
according to the palaeogeographic reconstructions proposed by
Steininger and Rögl (1985) and Meulenkamp and Sissingh
(2003), during this time interval a connection between palaeo-
Mediterranean and Paratethys probably still existed through the
present Marmara Sea area. The ecological barrier that existed
between the two palaeobioprovinces (marine realm in the
palaeo-Mediterranean and brackish conditions in the Para-
tethys) prevented any faunal exchange among them.
The late Messinian was characterized by the closure of the
palaeo-Mediterranean/Atlantic connection through the Bethic
and Riff Straits (Hsü et al., 1973; Cita, 1973; Benson, 1975;
Weijermars, 1988), which completely changed the palaeogeo-
graphic and palaeoenvironmental conditions of the Mediterra-
nean area. After the evaporite deposition due to the Messinian
Salinity Crisis, the palaeo-Mediterranean was affected by a
severe humid climate phase (Zeit Wet Phase sensu Griffin,
2002) linked to astronomical cyclicity (Hilgen, 1991; Krijgs-
man et al., 1999) and the palaeo-Mediterranean waters became
diluted down to oligo-mesohaline salinities. This caused the
disruption of the ecological barrier between palaeo-Mediterra-
nean and Paratethys and the westward migration of the
Paratethyan fauna that entered and flourished in the palaeo-
Mediterranean whose local fauna had been severely impover-
ished by the salinity crisis and the following water dilution.
Finally, at the beginning of the Pliocene, the re-establishment of
the Atlantic-palaeo-Mediterranean connection, due to the

Fig. 1. Structural sketch map of the Mediterranean and Paratethys (ruled areas), with the location of the late Messinian Lago-Mare localities mentioned in Table 7. 1. Vera
Basin (Spain) Carbonnel, 1978a; Cita et al., 1980; Benson and Rakic-El Bied, 1991). 2. Sorbas Basin (Spain) (Roep and Th. Van Harten, 1979; Krstić and Stancheva,
1990). 3. ODP 978 (Alboran Basin) (Iaccarino and Bossio, 1999). 4. DSDP 372 (Balearic Basin) (Benson, 1978). 5. ODP 975 (Balearic Basin) (Iaccarino and Bossio,
1999). 6. Rhone Basin (France) (Carbonnel, 1978a). 7. Aleria Basin (Corse, France) (Carbonnel, 1978a). 8. Caprera Canyon (Tyrrhenian Basin) (Aleria, 1979). 9. ODP 974
(Tyrrhenian Basin) (Iaccarino and Bossio, 1999). 10. ODP 654 (Tyrrhenian Basin) (Cita et al., 1990). 11. ODP 652 (Tyrrhenian Basin) (Borsetti et al., 1990). 12.
Castell’Arquato, Casa Giovanni Bello, Savignano sul Panaro (Italy). 13. Monticino quarry, Cella (Italy). 14. Perticara, Sapigno, Agip drill-holes, Buttafuoco (Italy). 15.
Pergola, Maccarone (Italy). 16. Cava Serredi, Valdelsa Basin, Volterra Basin, Ribolla Basin, Casino Basin, Podere S. Pace, Torrente Morra (Italy). 17. Blera, Civitella Cesi
(Italy). 18. Mondragone well (Italy). 19. Le Vicenne (Italy). 20. Roccacaramanico, Roccamorice, S. Valentino in Abruzzo Citeriore, Ca’Ciucco, Fonte dei Pulcini (Italy).
21. Eraclea Minoa (Italy). 22. Pasquasia (Italy). 23. Strimon Basin (Greece) (Gramann, 1969). 24. Aegina Island (Greece) (Krstić and Stancheva, 1990). 25. Corfou
(Greece) (Vismara-Schilling et al., 1976). 26. Crete (Greece) (Cosentino et al., 2007 this volume). 27. Kos Island (Greece) (Guernet et al., 1976). 28. DSDP 375 (Eastern
Mediterranean) (Benson, 1978). 29. DSDP 376 (Eastern Mediterranean) (Benson, 1978). 30. ODP 968 (Eastern Mediterranean) (Rouchy et al., 2001).
Fig. 1. Carte structurale de la Méditerranée et de la Paratéthys avec la localisation géographique des gisements du Lago Mare (Messinien supérieur) mentionnés dans
le Tableau 7.
 E. Gliozzi et al. / Geobios 40 (2007) 325–337
extensional tectonics, which affected the Gibraltar area,
restored fully marine conditions to the palaeo-Mediterranean
(Zanclean deluge, Benson, 1986).
Miocene ostracod palaeobiogeography seems to match well
the palaeogeographic reconstructions of Middle and Late
Miocene. Jiřiček and Řı́ha (1991) show that ostracods record
the latter marine connection between palaeo-Mediterranean
and Paratethys during the Serravallian (Badenian), with the
ostracod Paratethyan zones Acanthocythereis histrix/Bythocy-
pris lucida, Eocytheropteron inflatum/Falunia spinulosa,
Pajenborchella laskarevi/Neocyprideis (Miocyprideis) sarma-
tica elongata and Carinocythereis carinata/Phlyctenophora
farkasi, while no palaeo-Mediterranean ostracods migrating
into the Paratethys domain are recognized during the Tortonian-
early Messinian or vice-versa. During this time interval the
Paratethyan ostracod assemblages show a strong endemic
character, while the palaeo-Mediterranean marine ones shows
strong similarities to the Atlantic ostracods (Benson, 1976,
1984, 1990). On the contrary, during the post-evaporitic late
Messinian, the palaeo-Mediterranean ostracod assemblages,
from Spain to Cyprus, are dominated by a rich contingent of
ostracods with Paratethyan affinity (late Messinian Lago-Mare
biofacies) (Gliozzi et al., 2002, 2005, and references therein).
Notwithstanding the documented palaeogeographic isola-
tion of palaeo-Mediterranean and Paratethys during the
Tortonian-early Messinian, within some Italian brackish
deposits of this period some ostracods with Paratethyan affinity
have been recovered. The aim of this paper is to detail such taxa
and to explain the different dispersal modes that affected
ostracods during the late Tortonian-early Messinian on one side
and late Messinian on the other.
2. Late Tortonian-early Messinian Paratethyan
immigrants
Late Tortonian-early Messinian Italian brackish deposits are
known mainly from Tuscany (central Italy) (Bossio et al., 2002
and references therein). Recently, several sampling pro-
grammes have been carried out for micropaleontological
analyses and, among a rich contingent of Italian endemic or
central European species, a few brackish and freshwater
ostracods with Paratethyan affinity have been recognised. In
particular, the authors have directly investigated three basins:
the Velona Basin (Ghetti et al., 2002), the Valdelsa Basin
(unpublished data) and the Cinigiano-Baccinello Basin
(Benvenuti et al., 2004). Further data has been obtained from
the literature (Devoto, 1968; Bossio et al., 1994a, 1994b, 1994c,
1996, 2002; Testa, 1995) concerning other Tuscan localities as
Belforte (Siena), Val di Cornia (Grosseto), Casino Basin and
Volterra Basin (Fig. 2). The Paratethyan genera recovered in
these deposits are Bakunella Schneider, 1958, Camptocypria
Zalányi, 1944, Labiatocandona Krstić and Stancheva, 1990,
Lineocypris Zalányi, 1929, Propontoniella Krstić, 1972 (all
included in the subfamily Candoninae), Loxoconchissa
(Loxoconchissa) Triebel and Malz, 1969 and Loxoconchissa
(Loxocaspia) Schornikov, 1973 (included in the family
Loxoconchidae) and Amnicythere Devoto, 1965, Chartocythere
327
Livental in Buryndina, 1969, and Mediocytherideis Mandel-
stam 1956 (included in family Leptocytheridae).
All of the mentioned Candoninae originated in the Late
Miocene of the Paratethys, except for Propontoniella, which,
according to Krstič and Shao-Zeng, 2000 probably appeared in
the Late Palaeogene of China. Camptocypria includes living
representatives in the Caspian lake (Naydina, 1970; Schorni-
kov, 1966; Boomer et al., 2005).
Loxoconchissa s.s. originated during the Maeotian (Torto-
nian) of Romania (Triebel and Malz, 1969) while Loxoconch-
issa (Loxocaspia), whose living representative [L. (Loxocaspia)
immodulata (Stepanajtis)] is known from the Aral Lake, is also
known as fossil from the Plio-Pleistocene of Paratethys and
Greece (Agalarova et al., 1961; Krstić and Dermitzakis, 1981;
Mostafawi, 1994). Thus, the late Tortonian and early Messinian
Italian species of L. (Loxocaspia) seems to represent the more
ancient forms of this taxon. At present, taking into account the
geographical distribution there is no evidence to suggest a
palaeo-Mediterranean origin for this genus.
Amnicythere, Chartocythere and Mediocytherideis are
probably Paratethyan genera distributed from the Sarmatian
(Serravallian) to the Pontian (Messinian) (Mediocytherideis up
to the Early Pleistocene, Amnicythere up to present with living
representatives in the Caspian and Black Seas) (Stancheva,
1968; Krstić and McKenzie, 1991). On examining some
Leptocytheridae specimens from the Volterra Basin (Radicon-
doli area), Gliozzi et al. (2005) excluded the presence in Italy of
Chartocythere and referred them to the palaeo-Mediterranean
subgenus Mediocytherideis (Sylvestra) Doruk. The possibility
to see other samples recorded in the Volterra Basin confirm the
presence of at least three or four species referable to
Chartocythere, while, at present, the authors of this paper
are not convinced of the validity of the subgenus Sylvestra,
which could be considered as a junior synonym of Medio-
cytherideis s.s. If further taxonomic studies will confirm this
hypothesis, the genus Mediocytherideis, can no longer be
considered a Paratethyan taxon.
In conclusion, apart from the Leptocytheridae, which gave
rise to a great adaptive radiation in the Paratethyan domain but
whose origins date to the Serravallian, a period when the
palaeo-Mediterranean and Paratethys were still in partial
connection, all the other genera appeared in the Paratethys after
the separation of the two realms; they can therefore be
considered true Paratethyan immigrants.
The Italian forms pertaining to the previously mentioned
genera collected in the late Tortonian-early Messinian of Italian
brackish basins are reported in Table 1 from which it can be
seen that during this time interval the Paratethyan affinity is
limited to a generic or subgeneric rank. In fact, almost all the
Paratethyan taxa gave rise, in Italy, to new, endemic species, or
are represented by few valves (often early instars) left in open
nomenclature.
3. Late Messinian Lago-Mare Paratethyan immigrants
The Italian late Messinian Lago-Mare deposits outcrop
widely along the whole peninsula and have been intensively
328 E. Gliozzi et al. / Geobios 40 (2007) 325–337

Fig. 2. Structural sketch map of Tuscany with the location of the studied late Tortonian-early Messinian basins.
Fig. 2. Carte structurale de la Toscane avec la localisation des basins étudiés du Tortonien supérieur-Messinien inférieur.

studied (Table 2). They are characterized by several rich brackish
ostracod assemblages made up by a dominant Paratethyan
contingent. Moreover, during this period the very strong affinity
between palaeo-Mediterranean and Paratethyan assemblages
can be detected at the specific level. In fact, apart from few
palaeo-Mediterranean endemic species such as Cyprideis
agrigentina, Cyprideis anlavauxensis, Cyprideis tessalonikae,
Cyprideis sp. five sensu Gliozzi and Grossi (2004) (which,
according to Krstić, 1968a, 1968b and Carbonnel, 1978b, 1980
are not related with the Cyprideis species of the Paratethyan
bioprovince), Cypria robusta and some Amnicythere species
still in open nomenclature, several Paratethyan species have also
been recovered (Table 3). At present, 68 species have been
recognized within the late Messinian Lago-Mare deposits: six
(10.3%) are represented by palaeo-Mediterranean species
referable to the genera Cyprideis and Cypria, 38 (55.9%) have
been identified as Paratethyan species while the remaining 24
(35.3%) are still in open nomenclature, but their affinity with
Paratethyan species has been remarked upon. According to
Bassetti et al. (2003) the importance of the Paratethyan
contingent within the palaeo-Mediterranean Lago-Mare ostra-
cod assemblages has been overestimated. They recognize the
presence of a high number of new species referable to
Paratethyan genera for which they suppose a palaeo-Mediterra-
nean speciation. This speciation should have occurred either
during the 200 Ka long Lago-Mare event or in a longer time
interval, deriving from yet unknown species, which inhabited the
palaeo-Mediterranean brackish waters before and during the
 E. Gliozzi et al. / Geobios 40 (2007) 325–337 329

Table 1
Italian late Tortonian-early Messinian taxa referred to Paratethyan genera
Tableau 1
Taxa italiens du Tortonien supérieur-Messinien inférieur se rapportant à des genres paratéthysiens
Species Basin
Bakunella sp. (juv.) Cinigiano-Baccinello Basin
Camptocypria sp. (juv.) Cinigiano-Baccinello Basin
Camptocypria sp. Volterra Basin
Camptocypria aff. C. ossoinae milonovici Krstic Marsiliana
Camptocypria aff. C. labiata Val di Cornia
Volterra Basin
Casino Basin
Marsiliana
Candona cf. C. postsarmatica Krstic Casino Basin
Labiatocandona globosa Ceci and Gliozzi Valdelsa Basin
Lineocypris sp. 1 (juv.) Valdelsa Basin
Lineocypris sp. 2 (juv.) Valdelsa Basin
Propontoniella sp. Cinigiano-Baccinello Basin
Loxoconchissa (Loxoconchissa) kinoi Faranda, Cinigiano-Baccinello Basin
Gliozzi and Ligios
Loxoconchissa (Loxocaspia) cosentinoi Faranda, Velona Basin
Gliozzi and Ligios
Loxoconchissa (Loxocaspia) nuda Faranda, Cinigiano-Baccinello Basin
Gliozzi and Ligios
Loxoconchissa (Loxocaspia) punctata Faranda, Velona Basin
Gliozzi and Ligios
Loxoconchissa (Loxocaspia) reticulata Faranda, Velona Basin
Gliozzi and Ligios
Loxoconchissa (Loxocaspia) tuberosa Faranda, Cinigiano-Baccinello Basin
Gliozzi and Ligios
Loxoconchissa (Loxocaspia) velonae Faranda, Velona Basin
Gliozzi and Ligios
Amnicythere palimpsesta (Livental). Casino Basin
Amnicythere aff. A. palimpsesta (Livental). Marsiliana
Amnicythere pequenita (Stancheva) Belforte
Amnicythere sp. Val di Cornia
Casino Basin
Chartocythere spp. (3 species) Val di Cornia
Volterra Basin
Marsiliana
Chartocythere cf. C. franzi Marsiliana
Mediocytherideis spp. (7 species) Velona Basin
Volterra Basin
Age References
early Messinian Benvenuti et al., 2004
late Tortonian Benvenuti et al., 2004
early Messinian Bossio et al., 1996
late Tortonian-early Messinian Bossio et al., 1994c
late Tortonian-early Messinian Bossio et al., 1994b;
Testa, 1995; Bossio et al., 1996
Bossio et al., 2002
Bossio et al., 1994c
late Tortonian-early Messinian Bossio et al., 2002
late Tortonian-early Messinan Unpublished data
late Tortonian-early Messinan Unpublished data
late Tortonian-early Messinan Unpublished data
early Messinian Benvenuti et al., 2004
early Messinian Faranda et al., 2007 this volume
early Messinian Faranda et al., 2007 this volume
middle Tortonian-early Messinian Faranda et al., 2007 this volume
early Messinian Faranda et al., 2007 this volume
early Messinian Faranda et al., 2007 this volume
middle Tortonian-early Messinian Faranda et al., 2007 this volume
early Messinian Faranda et al., 2007 this volume
late Tortonian-early Messinian Bossio et al., 2002
late Tortonian-early Messinian Bossio et al., 1994c
early Messinian Devoto, 1968
late Tortonian-early Messinian Bossio et al., 1994b;
Bossio et al., 2002
late Tortonian-early Messinian Bossio et al., 1994b;
Testa, 1995
Bossio et al., 1994c
late Tortonian-early Messinian Bossio et al., 1994c
early Messinian Ghetti et al., 2002
late Tortonian-early Messinian Bossio et al., 1994a

salinity crisis. We partially disagree with these hypotheses since
we recognize a high similarity at a specific level with the Pontian
Paratethyan ostracods. We think that to the 54% of Paratethyan
species discussed above it is reasonable to add also the 35%
related to the Paratethyan-like taxa. In fact, sometimes
difficulties in identifying the valves at specific level is linked
to several causes such as bad preservation, scarcity of material,
unclear illustrations of the type-specimens published within old
Russian papers. Often slight differences of dimensions or
ornamentation have been overvalued, without taking into
account that these species lived in unstable water environments
characterized by salinity fluctuations that could be responsible
for ecophenotypical changes within the same taxon. Anadon
et al. (2002) and Boomer and Eisenhauer (2002) report several
examples of ecophenotypical characters recognized in marginal
marine ostracod species included in genera Cyprideis, Lox-
oconcha, Leptocythere, Hemicyprideis, Cytheromorpha, due to
temperature and salinity variations. We agree with Bassetti et al.
(2003) that limited palaeo-Mediterranean speciation events
could have occurred, giving rise to new subspecies or species, but
we think that their hypothesis of speciation during the Lago-Mare
event is more reliable because, as illustrated in the previous
chapter, the pre-evaporitic Late Miocene brackish ostracod
assemblages of Italy show only a few Paratethyan genera, often
represented only by juvenile valves. It is our opinion that the
Paratethyan affinity of the Italian Lago-Mare assemblages could
reach over 90% of species.
4. Discussion
In order to evaluate the degree of affinity between the Late
Miocene Italian and Paratethyan brackish ostracod assem-
blages three tables have been prepared (Tables 4–6) in which
the genera distributed in late Tortonian (Late Pannonian), early
Messinian (early and middle Pontian) and Late Messinian (Late
Pontian) in Italy and Paratethys are listed. To better understand
330 E. Gliozzi et al. / Geobios 40 (2007) 325–337

Table 2
Italian late Messinian Lago-Mare localities
Tableau 2
Localités italiennes du Lago Mare (Messinien supérieur)
Localities Regions References
Castell’Arquato Emilia Grekoff and Molinari, 1963
Casa Giovanni Bello Emilia Krstić and Stancheva, 1990
Savignano sul Panaro Emilia Fregni et al., 2003
Monticino quarry Romagna Colalongo, 1988
Cella Romagna Colalongo et al., 1978
Sapigno Marche Bassetti et al., 2003
Perticara Marche Gliozzi and Grossi, 2004
AGIP Drill-holes (Montepetra, Ca’Blindana) Marche Unpublished data
Buttafuoco Marche Unpublished data
Pergola Marche Molinari Paganelli, 1975
Maccarone Marche Carbonnel, 1978a
Ca Ciucco Abruzzo Carbonnel, 1978a
Roccamorice, Roccacaramanico Abruzzo Patacca et al., 1992
S. Valentino in Abruzzo Citeriore Abruzzo Cipollari et al., 1999b, Gliozzi et al., 2002
Le Vicenne Abruzzo Colacicchi et al., 1967, Cipollari et al., 1999a, Gliozzi, 1999.
Fonte dei Pulcini Abruzzo Cipollari et al., 2003
Valdelsa Basin Tuscany Bossio et al., 1993b
Ribolla Basin Tuscany Bossio et al., 1993a
Volterra basin (several localities) Tuscany Bossio et al., 1994a, 1996
Podere S. Pace (Massa Marittima) Tuscany Bossio et al., 1976
Torrente Morra (Leghorn) Tuscany Bossio et al., 1997
Cava Serredi Tuscany Bossio et al., 1981
Casino Basin Tuscany Bossio et al., 2002
Blera, Civitella Cesi Latium Gliozzi et al., 2002
Mondragone well Campania Cipollari et al., 1999b
Eraclea Minoa, Capo Rossello Sicily Bonaduce and Sgarrella, 1999
Pasquasia-Capodarso Sicily Colalongo, 1967
ODP 974 Tyrrhenian Sea Iaccarino and Bossio, 1999
ODP 654 Tyrrhenian Sea Cita et al., 1990
ODP 652 Tyrrhenian Sea Borsetti et al., 1990

the assemblage composition for each period, the number of
species for each genus has been reported. Data for Paratethyan
ostracod assemblages are taken mainly from the results of the
International Geological Correlation Program ‘‘Stratigraphic
correlation Tethys-Paratethys Neogene’’, (Jiřiček, 1985; Krstić,
1985; Sokač, 1985; Krstić and Stancheva, 1990, Marinescu
et al., 1990; Olteanu, 1990, 1995; Sokač, 1990a, 1990b), to
which other specific papers have been added (Agalarova et al.,
1940, 1961; Agalarova, 1967; Sokač, 1972; Olteanu and Vekua,
1989; Stancheva, 1989, 1990; Pipı́k, 1998, 2001; Tonoğlu and
Gökçen, 1997; Tonoğlu, 2002). Ostracods from the Caspian-
Aral basin have not been reported, since this basin is very far
from the palaeo-Mediterranean and its palaeontological litera-
ture, all in Russian, is rather old, thus not taxonomically
updated. In Table 4 (Pannonian/Maeotian/Tortonian) the
Euxinic basin has not been included since during this time-
interval, according to the palaeogeographic reconstructions by
Hámor (1988) and Steininger and Rögl (1985), this basin was
still characterized by a polyhaline environment. This recon-
struction is confirmed by the Maeotian ostracod assemblages
recovered by Tonoğlu and Gökçen (1997), which show a clear
marine affinity.
It is possible to see from Tables 4–6, that during late
Tortonian and early Messinian very few Paratethyan genera,
which lived in the Pannonian, Dacic and Euxinic Basins, have
been recovered in Italy (Tables 4 and 5) while, during the late
Messinian Lago-Mare event almost all the Paratethyan genera
that populated the Pontian Paratethyan waters are present in the
Italian assemblages (Table 6).
If we consider some selected pre-evaporitic Late Miocene
deposits, whose ostracod assemblages have been investigated in
detail such as the Velona Basin, Cinigiano-Baccinello Basin
and Valdelsa Basin (Ghetti et al., 2002; Benvenuti et al., 2004;
unpublished data) the percentages of palaeo-Mediterranean
taxa and Paratethyan-like taxa have been calculated (Table 7),
which range 57–69% and 30–42%, respectively. These
percentages are far from those calculated for the late Messinian
ostracod assemblages (10% palaeo-Mediterranean taxa, 35%
Paratethyan-like taxa, 54% Paratethyan species). The differ-
ences are much more striking if we take into account that a high
percentage of late Tortonian-early Messinian Paratethyan-like
taxa are referred to genera which are not represented in the
Italian Lago-Mare ostracod assemblages (Mediocytherideis, L.
(Loxocaspia), Bakunella and Propontoniella). Similar percen-
tages in the ostracod assemblage composition have also been
calculated from Late Miocene West palaeo-Mediterranean
localities such as the brackish deposits of France (Carbonnel,
1969, 1978a) and Spain (Carbonnel, 1978a; Gliozzi et al., 2005)
(Table 7), showing that the composition of Late Miocene
ostracod assemblages in the palaeo-Mediterranean is rather
 E. Gliozzi et al. / Geobios 40 (2007) 325–337
Table 3
Italian late Messinian Lago-Mare species with Paratethyan affinity at a specific or generic level.
Tableau 3
Espèces italiennes du Lago Mare (Messinien supérieur) d’affinité paratéthysienne
Amnicythere anormalis (Suzin)
Amnicythere cellula (Livental)
Amnicythere costata (Olteanu)
Amnicythere idonea Mandelstam, Markova, Rozyeva and Stepanaytis/
A. pontica (Scheydaeva)
Amnicythere ex gr. A. larga Krstić
Amnicythere lata (Schneider)
Amnicythere litica (Livental)
Amnicythere multituberculata (Livental)
Amnicythere palimpsesta (Livental)
Amnicythere liticoides Krstić and Stancheva
Amnicythere propinqua (Livental)
Amnicythere aff. A. propinqua (Livental) (Miculan in Bassetti et al., 2004)
Amnicythere aff. A. resupina (Bonaduce and Sgarrella, 1999)
Amnicythere rosalinae (Schneider in Agalarova, 1940)
Amnicythere soluta (Livental)
Amnicythere subcaspia (Livental in Agalarova et al., 1940)
Amnicythere sp. 1 (Gliozzi and Grossi, 2004)
Amnicythere sp. 1 (Gliozzi et al., 2002)
Amnicythere sp. 2 (Gliozzi et al., 2002)
Amnicythere sp. C (Miculan in Bassetti et al., 2004)
Amnicythere sp. D (Miculan in Bassetti et al., 2004)
Amnicythere sp. G (Miculan in Bassetti, 2004)
Camptocypria aff. C. glabra (Krstić)
Camptocypria sp. 1 (Gliozzi and Grossi, 2004)
Candona cf. C. postsarmatica Krstić
Caspiocypris pontica (Sokač)
Caspiocypris cf. C. alta (Zalányi)
Chartocythere sp. (Bossio et al., 1996, 1986)
Cypria robusta (Devoto)
Cyprideis agrigentina Decima
Cyprideis anlavauxensis Carbonnel
Cyprideis ex gr. torosa (Jones) (Bossio et al., 1986)
Cyprideis thessalonikae Krstić
Cyprideis sp. 5 (Gliozzi and Grossi, 2004)
homogeneous, suggesting that their distribution is probably
driven by the palaeogeography. Further confirmation of this
hypothesis come indirectly from the composition of the late
Serravallian ostracod assemblages recovered in Spain (Ebro
Basin, Gliozzi et al., 2005) (Table 7). While the percentage of
palaeo-Mediterranean and Paratethyan-like taxa are rather
comparable with the Tortonian-early Messinian ones, in the late
Serravallian assemblages a few Paratethyan species are still
present (23.5%) whose presence is probably linked to the
northern connection with the Paratethyan domain which was
closing just at that time (Meulenkamp and Sissingh, 2003).
In conclusion, the data suggest that the Italian late Tortonian-
early Messinian and late Messinian Lago-Mare brackish
ostracod assemblages are characterized by a different degree
of affinity with the Paratethys bioprovince: in the first case the
Paratethyan contingent is smaller and the affinity is limited to
the generic or subgeneric level while in the second case the
Paratethyan contingent made up the majority of the assemblages
and the affinity reaches the specific level. Such differences in
the palaeobiogeographical composition of the Late Miocene
ostracod assemblages match the different palaeogeographic
331
Cyprideis ex gr. tuberculata Mehés (Bossio et al., 1986; Gliozzi, 1999)
Cytherura pyrama Schneider
Euxinocythere (Euxinocythere) suzini (Scheneider)
Euxinocythere (Maeotocythere) bosqueti (Livental)
Euxinocythere (Maeotocythere) aff. E. (M.) bosqueti (Livental)
Euxinocythere (Maeotocythere) aff. E. (M.) moesica Olteanu
Euxinocythere (Maeotocythere) praebaquana (Livental in Agalarova et al., 1940)
Euxinocythere (subgen. ?) lacunosa (Reuss)
Labiatocandona aff. L. labiata (Zalányi)
Labiatocandona aff. C. portaferrica Pokorný and Stancheva, 1990
Lineocypris cf. L. fossulata (Pokorný)
Lineocypris cf. L. hodonensis (Pokorný)
Lineocypris sp. 1 (Gliozzi and Grossi, 2004)
Lineocypris sp. (Molinari Paganelli, 1975)
Lineocypris sp. (Carbonnel, 1978a)
Loxoconcha ancilla Stancheva
Loxoconcha eichwaldi Livental
Loxoconcha aff. L. kochi Mehés
Loxoconcha cf. L. ludica Olteanu
Loxoconcha muelleri Mehés
Loxoconcha rhombovalis Pokorný
Loxoconcha cf. L. schweyeri dacica Olteanu
Loxocorniculina djafarovi (Schneider in Suzin)
Mediocytherideis (Sylvestra) sp. (Miculan in Bassetti et al., 2004)
Pontoniella pontica (Agalarova) n. ssp. (Gliozzi and Grossi, 2004)
Pontoniella aff. P. pontica (Agalarova) (Bonaduce and Sgarrella, 1999)
Pontoniella sp. (Molinari Paganelli, 1975)
Pseudocythere limata (Schneider in Agalarova et al., 1940)
Semicytherura aff. S. inversa (Seguenza)
Typhlocypris cf. T. redunca (Zalányi)
Tyrrhenocythere ruggierii Devoto
Tyrrhenocythere pontica (Livental)
Tyrrhenocythere cf. T. taurica Sinegub in Krstić, 1977
Zalanyiella venusta (Zalányi)
settings of the palaeo-Mediterranean-Paratethys area and they
must reflect different dispersion modes.
The geographic occurrence of the late Tortonian-early
Messinian Paratethyan-like ostracods shows a disjunct dis-
tribution, which is consistent with a passive dispersal. Griffiths
and Holmes (2000) summarize the possible passive dispersal
modes of freshwater ostracods, which include wind, aquatic
birds, aquatic insects, amphibians, fishes and man. Wind and
aquatic insects would seem to be dispersal agents only for short
distances (Thienemann, 1950; Fryer, 1953) and, in the case of
Paratethys/palaeo-Mediterranean relations, amphibians and
fishes must be rejected since there was no aquatic continuity
between the two domains. Thus, the most reliable passive
dispersal agent seems to be aquatic birds. Many papers
document this dispersal mode; freshwater ostracods are proved
to have been transported either as individuals or as eggs directly
or through mud attached to bird bodies (feet, feathers and
beaks) (De Deckker, 1977; Löffler, 1964; Horne and Smith,
2004). Several papers showed that Cypridinid eggs preserve
their vitality during the passage through the digestive tract of
aquatic birds (Proctor, 1964; Proctor and Malone, 1965;
332 E. Gliozzi et al. / Geobios 40 (2007) 325–337

Table 4
Late Tortonian ostracod genera distributed in Italy and in the Paratethyan domains
Tableau 4
Genres d’ostracodes du Tortonien supérieur distribués en Italie et dans les domaines paratéthysiens
Italian basins Pannonian basin Dacic Basin
Amnicythere 8 Amnicythere 4
Amplocypris 16
Bakunella 1
Bononiella 1 Bononiella 6
Camptocypria 3 Camptocypria 11 Camptocypria 3
Chartocythere 4 Chartocythere 1
Caspiocypris 1 Caspiocypris 1
E.Euxinocythere) 1 E. (Euxinocythere) 1
E.(Maeotocythere) 1 E. (Maeotocythere) 11
H. (Hemicytheria) 12
H. (Getocytheria) 16
Hungarocypris 4
Labiatocandona 1
Lineocypris 6 Lineocypris 1
Loxocorniculina 1
L. (Loxocaspia) 9
L.(Loxoconchissa) 2
Mediocytherideis 3 Mediocytherideis 1
Ochridella 3
Pontocythere 1
Pontoleberis 3 Pontoleberis 1
Pontoniella 2 Pontoniella 3
Propontoniella 1
Reticulocandona 2
Serbiella 7
Sinegubiella 2
Stanchevia 2 Stanchevia 4
Thaminocypris 3
Turkmenella 1 Turkmenella? 4
Typhlocyprella 3
Typhlocypris 2 Typhlocypris 1
Tyrrhenocythere 6
Zalanyiella 2 Zalanyiella 3

McKenzie, 1971). Thus, we can conclude that the occasional
dispersion of Paratethyan-like taxa during late Tortonian-early
Messinian probably occurred through aquatic birds. If it is true
what stated by Morales (1993) that the migratory behaviour of
birds seems to appear later (Early Pleistocene) we must suppose
the existence of aquatic birds whose areal distribution was
extended E-W from Paratethys to palaeo-Mediterranean.
On the contrary, when late Messinian Lago-Mare ostracod
assemblages are considered, the areal distribution of the
Paratethyan species seems to be continuous, from the
Dardanelli area to the west (Gliozzi et al., 2002 and references
therein). At that time, apart from some small thrust-top brackish
basins, the palaeo-Mediterranean was characterized by several
huge brackish basins partially connected to each other (Benson,
1986; McCulloch and De Deckker, 1989), thus the Paratethyan
ostracod dispersion could have been performed both by
hydrological pathways and by passive dispersal via aquatic
birds. In both cases the dispersal rate and the efficiency of the
dispersion were higher than in the late Tortonian-early
Messinian, due to more favourable palaeogeographic condi-
tions, giving rise to a complete ostracod assemblage turnover in
only a few tens of thousands of years. Paratethyan species that
lived during the Late Pontian in the eastern Pannonian, Dacic
and western Euxinic Basins were more diversified than the
Paratethyan species, which effectively entered the palaeo-
Mediterranean. In particular the major differences concern the
pointed candonids, clustered in this informal group for their
pointed postero-ventral border, referable to genera Camptocy-
pria, Pontoniella, Lineocypris and Zalanyiella. Only two
species (P. pontica and Z. venusta) against 38 potential
immigrants has been recognized in the late Messinian Italian
Lago-Mare deposits. Surely this figure is slightly under-
estimated, due to identification problems, but, even if we take
into account the Italian pointed candonids left in open
nomenclature, their numbers rise to a maximum of only ten
species (Table 3). According to some palaeoecological data, the
Messinian Lago-Mare pointed candonids seem to be char-
acteristic of oligo- to mesohaline rather deep waters (Gofman,
1966; Gliozzi and Grossi, 2004). On the contrary, shallow
mesohaline genera such as Amnicythere, E. (Maeotocythere),
Loxocorniculina and Loxoconcha are represented in the Italian
late Messinian by the 54% of the potential Paratethyan
immigrants. These figures could be explained if it were
assumed that shallow mesohaline environments were more
 E. Gliozzi et al. / Geobios 40 (2007) 325–337 333

Table 5
Early Messinian ostracod genera distributed in Italy and in the Paratethyan domains
Tableau 5
Genres d’ostracodes du Messinien inférieur distribués en Italie et dans les domaines paratéthysiens
Italian basins Pannonian basin Dacic basin Euxinic basin
Amnicythere 4 Amnicythere 9 Amnicythere 23 Amnicythere 5
Amplocypris 9 Amplocypris 1 Amplocypris 1
Bakunella 1 Bakunella 2 Bakunella 3 Bakunella 1
Bononiella 1
Camptocypria 3 Camptocypria 9 Camptocypria 19 Camptocypria 4
Chartocythere 4 Chartocythere 1
Caspiocypris 4 Caspiocypris 2
Cryptocyprideis 1
E. (Euxinocythere) 1 E. (Euxinocythere) 2
E. (Maeotocythere) 2 E. (Maeotocythere) 3 E. (Maeotocythere) 2
Hastacandona 1 Hastacandona 3
H. (Hemicytheria) 8 H.(Hemicytheria) 2 H.(Hemicytheria) 2
Hungarocypris 2 Hungarocypris 1
Kowalskiella 1
Labiatocandona 1 Labiatocandona 3 Labiatocandona 1
Lineocypris 6 Lineocypris 1
Loxoconcha 1
L.(Loxoconchissa) 1
Loxocorniculina 1 Loxocorniculina 1
Mediocytherideis 7 Mediocytherideis 3
Pontoleberis 1 Pontoleberis 2
Pontoniella 5 Pontoniella 15 Pontoniella 1
Propontoniella 1
Pseudocythere 1
Reticulocandona 1
Serbiella 4 Serbiella 1 Serbiella 1
Sinegubiella 1
Thaminocypris 2
Typhlocyprella 1
Typhlocypris 1 Typhlocypris 1
Tyrrhenocythere 1 Tyrrhenocythere 8 Tyrrhenocythere 1
Zalanyiella 3 Zalanyiella 3

Table 6
Late Messinian Lago-Mare ostracod genera distributed in Italy and in the Paratethyan domains
Tableau 6
Genres d’ostracodes du Lago Mare (Messinien supérieur) distribués en Italie et dans les domaines paratéthysiens
Italian basins Pannonian basin Dacic basin Euxinic basin
Amnicythere 23 Amnicythere 8 Amnicythere 17 Amnicythere 6
Amplocypris 2 Amplocypris 1
Bakunella 4 Bakunella 1 Bakunella 1
Camptocypria 2 Camptocypria 11 Camptocypria 9 Camptocypria 2
Candona 1 Candona 1
Caspiocypris 2 Caspiocypris 4 Caspiocypris 3
Chartocythere 1
Cryptocyprideis 1
Cyprinotus 1
Cytherura 1
Dacicandona 1
E. (Euxinocythere) 1 E. (Euxinocythere) 1 E. (Euxinocythere) 2
E. (Maeotocythere) 4 E. (Maeotocythere) 2 E. (Maeotocythere) 4 E. (Maeotocythere) 2
Hastacandona 2
H. (Hemicytheria) 2
Hungarocypris 1
Labiatocandona 2 Labiatocandona 1 Labiatocandona 1
Lineocypris 5 Lineocypris 6 Lineocypris 3
Loxoconcha 8 Loxoconcha 4 Loxoconcha 5 Loxoconcha 1
Loxocorniculina 1 Loxocorniculina 1 Loxocorniculina 1
Mediocytherideis 1
Pontoleberis 1
334 E. Gliozzi et al. / Geobios 40 (2007) 325–337
Table 6 (Continued )
Italian basins Pannonian basin Dacic basin Euxinic basin
Pontoniella 3 Pontoniella 5 Pontoniella 6 Pontoniella 1
Pseudocythere 1 Pseudocythere 1
Reticulocandona 1
Semicytherura 1
Serbiella 1
Typhlocyprella 1
Typhlocypris 1 Typhlocypris 1
Tyrrhenocythere 3 Tyrrhenocythere 1 Tyrrhenocythere 6
Zalanyiella 1 Zalanyiella 2

Table 7
Comparisons between the percentage composition of the ostracod assemblages from Late Miocene in selected palaeo-Mediterranean localities (see Fig. 1)
Tableau 7
Comparaisons des compositions en % des associations d’ostracodes du Miocène supérieur dans les localités paléo-méditerranéennes les plus importantes (voir Fig. 1)
Basins Palaeo-Mediterranean Palaeo-Mediterranean species Paratethyan
taxa (%) with Paratethyan affinities (%) species (%)
Ebro Basin (late Serravallian) 52.9 23.5
Cinigiano- Baccinello (middle-late Tortonian) 62.5 37.5
Valdelsa Basin (late Tortonian-early Messinian) 69.2 30.8
Rhône Basin (Tortonian-early Messinian) 88.5 11.5
Velona Basin (early Messinian) 57.1 42.9
Cinigiano-Baccinello (early Messinian) 68.4 31.6
Several Italian Lago-Mare assemblages 10.3 35.3 54.4
Several French Lago-Mare assemblages 7.7 11.5 80.8
Several Spanish Lago-Mare assemblages 11.8 29.4 58.8

widely dispersed in the palaeo-Mediterranean and that the
Paratethyan species, which were able to enter the palaeo-
Mediterranean, were the most euryplastic forms among those
that made up the Paratethyan assemblages, thus more easily
adaptable to slightly different environments.
5. Conclusions
 notwithstanding the favourable palaeogeographic conditions
for the active dispersion during the late Messinian Lago-Mare
event, the palaeo-Mediterranean was colonized only by half
of the potential Paratethyan ostracod immigrants. It is
probable that the species that were able to enter the palaeo-
Mediterranean were the most euryplastic.

The analysis of the Italian brackish ostracod assemblages Acknowledgments

recovered in late Tortonian, early Messinian and late Messinian
Lago-Mare basins has shown that:
 during the late Tortonian and early Messinian the ostracod
Paratethyan affinity is recognizable only at a generic level.
This matches well with the palaeogeographic knowledge of
this period with the palaeo-Mediterranean separated from the
Paratethys (Meulenkamp and Sissingh, 2003). It is possible
that the few Paratethyan-like ostracods recovered in Italy,
may have followed a passive dispersal via aquatic birds
followed by endemic Italian speciation;
 during the late Messinian Lago-Mare event, the Paratethyan
affinity of the Italian ostracod assemblages rises to the
specific level, following the new palaeogeographic condi-
tions of the Atlantic/palaeo-Mediterranean/Paratethys
domains (palaeo-Mediterranean/Atlantic isolation, palaeo-
Mediterranean/Paratethys connection). During this period an
active dispersal mode through the more or less continuous
waterbodies can be supposed, together with a more efficient
passive dispersal via aquatic birds. Palaeo-Mediterranean
speciation events were limited;
We wish to thank Ian Boomer and Radovan Pipı́k for their
valuable suggestions, which helped us to improve this paper.
This work was financially supported by MIUR (Cofin 2003,
Resp. D. Cosentino).
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