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Evolution of late Miocene mammalian faunas and palaeoenvironment in the Northern and Eastern Mediterranean realms.

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5
th International Symposium on Eastern Mediterranean Geology
Thessaloniki, Greece, 14-20 April 2004
Evolution of late Miocene mammalian faunas
and palaeoenvironment in the Northern and
Eastern Mediterranean realms
Bonis L. de1, Bouvrain G.2 & Koufos G.D.3
1Laboratoire de Geobiologie, Biochronologie et Paléontologie Humaine. Université de
Poitiers, 40 av. du rect. Pineau, F-86022, Poitiers, France, louis.debonis@univ-poitiers.fr
2Laboratoire de Paléontologie, Université Pierre et Marie Curie (Paris 6), 4 place Jussieu,
F-75005, Paris, France
3Laboratory of Geology and Palaeontology, Department of Geology, University of
Thessaloniki, GR-54124, Thessaloniki, Greece, koufos@geo.auth.gr
Keywords: Mammals, late Miocene, Mediterranean, palaeoecology
Introduction
The late Miocene is a crucial period in the history of peri-Mediterranean mammalian faunas
with a large turn-over and appearance of several new taxa in Europe. These change occurred
together with geological and geographical ones, as well as modification of climatic conditions.
Due to the movements of African and Eurasian plates, the compressive phase of the Alpine
orogenic procession, which begun during the Mesozoic, was active during the late Miocene.
This process determines the elevation of the mountains and the sliding up of slices, and also
a contraction of the area allowing faunal displacement, despite extension in some places.
This time corresponds to the finishing of the Appennine chain, the Sardo-Corsica area, the
Betic chain or the Kabylian mountains. During the Miocene a transgression invaded the peri-
Alps groove and gave another sea named Paratethys which spread from the Western Medi-
terranean to the Aral Sea and far ahead. From the late Miocene the Paratethys regressed
place to place to become sometimes lakes or lagoons and the spreading of the continental
areas allows some inter-continental relationships with the arrival of new faunas in Europe.
The continental late Miocene European mammalian faunas have been divided in two different
ages of mammals: Vallesian on the bottom and Turolian on the top (Crusafont 1954). The
Vallesian corresponds also to the mammalian Neogene zones MN 9 and MN 10 and the
Turolian to MN 11, MN 12 and MN 13 (Mein 1990). These ages of mammals were defined on
local Spanish faunas and the correlation with other areas in Europe sometimes is a problem.
Another problem will be the correlation with the Miocene marine stages In western Europe,
we admit that the Vallesian began a little after (11.3 ± 0.2 Ma) the Tortonian beginning and it
ended during the middle Tortonian (9.1 Ma). The Turolian corresponds to the upper Tortonian
and Messinian (9 to 5.5 Ma). In Central Paratethys, the former corresponds to the early to
middle Pannonian and the later to the late Pannonian and Pontian. In Eastern Paratethys, on
one hand, the Vallesian is correlated to the latest Sarmatian (Khersonian and may be late
Bessarabian) and the lower Maeotian and, on the other hand, the Turolian corresponds to the
late Maeotian and Pontian (Steiniger 1999).
Our aim is to follow the mammalian change through Europe first during the Vallesian and
second during the Turolian from the Iberian Peninsula to the Eastern Mediterranean. For
every age of mammals we shall look for faunal differences between the different areas and,
through the flora when possible (even if sometimes it is difficult to precisely correlate floristic
and mammalian localities), to explain the differences in terms of climatic variations. We statis-
tically compare the different large mammal genera associations to show that some of them
can be characteristic of climatic conditions (Bonis et al. 1979; Bonis et al. 1991; 1992).
Ref: S5-11
5
th International Symposium on Eastern Mediterranean Geology
Thessaloniki, Greece, 14-20 April 2004
Vallesian
For the Vallesian, a factor analysis founded on the per-cent (percentage) of localities in which
there are occurrences of a genus with each other genera reaches the result that there is a fau-
nal shift from east to west in the faunal associations, some genera could be linked with climatic
conditions. Another factor analysis comparing the localities themselves in function of their faunal
list shows that there are two different faunal sets in western and south eastern Europe corre-
sponding to a more forested and more open environment respectively. When mammals are as-
sociated with floral data, we can see that there is a floral change in Iberian Peninsula between
early Vallesian (MN 9) and late Vallesian (MN 10). This change corresponds to the "mid-
Vallesian crisis" (Sanz de Siria 1997; Agusti et al. 2003). In France, Switzerland, Germany and
central Europe (Austria, Hungary, Bohemia, Slovakia), the Vallesian faunas seems also linked
to quite forested areas (Bonis et al. 1991). The landscapes were very different in the Balkans
and southern Europe with the presence of open environment faunas. But in these areas we
cannot find a "mid-Vallesian crisis" because there are few localities or because there is no sig-
nificant change in the climatic conditions.
Turolian
The mammalian Turolian faunas as a whole are generally considered as open environment
faunas with few exceptions (Frantzen 1997: Frantzen & Storch 1975). But, an analysis of the
per-cent (percentage) localities in which a mammal genus is linked with another genus shows
that some genera are very often linked each other and it is possible to distinguish two groups,
one of them more associated to the presence of permanent water and probably less open envi-
ronment. The first group contains Castoridae, Tapiridae and Tragulidae which nowadays are
linked to the presence of water, and also Cervidae, Deinotheriidae, Chalicotheriidae , Dihoplus
and Dystichoceras. The other one contains Orycteropus which nowadays is ubiquistic and
Ceratotherium which nowadays deals with open environment. Both are grouped with Samoth-
erium, Chilotherium and Schizotheriinae, the last one being hypsodont and probably a grazer.
Another analysis (presence-absence in the Turolian localities) using only the mammalian gen-
era of both extreme groups on the graph allows to show what is the grouping of the localities.
2D Plot of Row Coordinates; Dimensions: 1 x 2
Input Table (Rows x Columns): 47 x 13
Standardization: Row and column profiles
FF
I
H
U
R
MM
G
E
T
EE
A
G
G
E
T
U
T
B
GTT
TT
E
E
T
FU
UU
EG
T
H
FY
E
T
SA SA
SA
T
U
G
E
-1,5 -1,0 -0,5 0,0 0,5 1,0 1,5 2,0
Dimension 1; Eigenvalue: ,65592 (29,28% of Inertia)
-2,5
-2,0
-1,5
-1,0
-0,5
0,0
0,5
1,0
1,5
Dimension 2; Eigenvalue: ,31019 (13,85% of Inertia)
Figure 1. Correspondence analysis : plotting of 47 fossil mammal localities on the
axis 1 and 2 in function of their faunal assemblages.
Ref: S5-11
5
th International Symposium on Eastern Mediterranean Geology
Thessaloniki, Greece, 14-20 April 2004
We note that the Western and Central European localities are opposed to the localities of
Samos and Turkey. The localities of continental Greece, Bulgaria and FYROM are in-between.
The localities of Moldavia spread from one group to another one on the top of the graph, those
of Ukraine are also on the top of the graph but close to the Greek and Bulgarian localities. The
analysis indicate differences in the paleo-environment of these areas as well as geographical
position. For instance, on one hand a German locality can plot close to the Spanish localities
insofar it contains the same kind of fauna corresponding to a similar environment. On the other
hand the grouping of the Samos localities with the Turkish ones could indicate the presence of
an Aegean strait during late Miocene. Mammals which characterise the Western and Central
European localities are more linked to humidity or forests and those which characterise the
eastern area seem to have dealt with more open environments.
Conclusions
The analysis of the late Miocene European mammalian faunas indicates that:
during Vallesian the palaeoenviromental conditions were more forested in Western
Europe and more open in South-eastern Europe.
during Turolian there two groups of mammalian faunas. The first is correlated with the
presence of permanent water and probably with less open environment, and the second
with more open environment and more drier conditions.
the Turolian localities of Western and Central Europe are opposite to those of Samos and
Turkey, while those of the Balkan Peninsula are between them.
the similarity of Samos and Turkish faunas indicates the possible presence of an Aegean
strait during the late Miocene.
References
Agusti J., Sanz de Siria A. & Garces M. (2003). Explaining the end of the hominoid experi-
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Bonis L. d., Bouvrain G. & Geraads D. (1979). Artiodactyles du Miocène supérieur de Macé-
doine, Annales géologiques des Pays Helléniques, 1, 167-175.
Bonis L. d., Bouvrain G., Geraads D. & Koufos G.D. (1991). Composition and species diversi-
ty in late Miocene faunal assemblages of northern Greece, Bulletin de la Société géo-
logique de Grèce, 25 (2), 395-404.
Bonis L. d., Bouvrain G., Geraads D. & Koufos G. D.(1992). Diversity and paleoecology of
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laeoecology, 91, 99-121.
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Annales de Paléontologie, 40, 97-103.
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Franzen J.L. & Storch G. (1975). Die unterpliozäne (turolische) Wirbeltierfauna von Dorn-
Dürkheim, Rheinhessen (SW-Deutschland). 1. Entdeckung, Geologie, Mammalia:
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(eds.), European Neogene Mammal Chronology, New York.
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lona), Paleontologia i Evolució, 30-31, 247-268.
Steininger F.F. (1999). Chronostratigraphy, Geochronology and Biochronology of the Miocene
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of Europe, Dr. Friedrich Pfeil, München.
Ref: S5-11
... Several scientists dealt with the determination of the Samos palaeoenvironment based on the whole mammal assemblage or on part of it (Solounias and Dawson-Saunders, 1988; Bonis et al., 1992a Bonis et al., , 1999 Bonis et al., , 2004 Solounias et al., 1999; Strömberg et al., 2007). In most of these analyses a key problem concerns the lack of adequate time resolution due to mixing of material from different horizons, representing different mammal associations and potentially different environments. ...
... The biogeographic relationships of the Samos mammal assemblages were discussed among palaeontologists and various aspects have been formulated (Bonis et al., 1992aBonis et al., , 2004 Bernor et al., 1996b; Solounias et al., 1999; Koufos et al., 2006). The two classical Turolian mammal faunas of Pikermi and Samos, although they are neighboring, differ significantly from a palaeoecological point of view, implying a subdivision of their common geographic domain [the so-called Greco- Iranian Province (Bonis et al., 1992b)] in smaller ecological clusters (Kostopoulos, 2009c). ...
Article
As one of the oldest known Eurasian fossil vertebrate localities, Samos late Miocene fauna attracted the interest of specialists by its richness and overall importance. Nevertheless, crucial taxonomical questions and chronological problems obscured its value. The detailed study of the local stratigraphy, the collection of new fossil material and its study, the revision of the old collections and the updated magneto-chronology of the fossiliferous deposits permited to re-discuss most of the problems in a special volume edited in 2009 by Koufos and Nagel and to provide a clearer and more precise idea about the Samos fauna and its age. A synopsis of this work is given here. The systematic study of the new collection (∼1200 identified specimens) allows the determination of 42 species from three fossil horizons, ranging from the upper part of early Turolian (MN11) to the end of middle Turolian (MN12). Taxonomic novelties are the presence of the carnivore genus Protictitherium found for the first time in Samos, the establishment of the new name Skoufotragus for Pachytragus Schlosser with the new species Skoufotragus zemalisorum, and the amended morphology of Pseudomeriones and Urmiatherium. Additionally six Hipparion and four Gazella species were recognized and a better morphometric distinction between Samotherium boissieri and Samotherium major was performed. This study also improved the correlation of the old fossiliferous sites with the new ones and with the local stratigraphy of the Mytilinii Basin, while precise ages have been obtained for the mammal localities. The new data together with the old collections indicate the presence in Samos of four chronologically successive mammal assemblages reflecting a “four stages-of-evolution” scheme. The Turolian palaeoenvironment of Samos is determined as an open bushland with thick grassy-floor of C3 graminoids with possible increase of the open and dry character from the beginning to the end of Middle Turolian. The Samos mammal faunas are palaeobiogeographically closer to the Asian ones than to those from the Greek mainland.
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The Vallesian Crisis involved the extinction of most of the hominoids that settled successfully in Europe during the middle and early Late Miocene, including Dryopithecus, Ankarapithecus and Graecopithecus. This event has been dated at 9.6 Ma, predating by more than one million years the spread of the C4 grasses and the retreat of forests over large parts of the globe at 7-8 Ma. The finding of macrofloral remains in the Terrassa section (Vallès-Penedès Basin) sheds new light on the nature of vegetational change associated with the hominoid extinction. This section presents an abundant late Vallesian vertebrate fauna and has been accurately dated at 9.2 - 9 Ma by paleomagnetism. Therefore, it provides the best indication of the kind of vegetation that occupied the area after the Vallesian Crisis. It is suggested that the extinction of the late Miocene Western European hominoids was not related to the spread of grasses, but to a significant increase of a floral association dominated by deciduous trees.
Quelques considérations biologiques sur le Miocène espagnol
  • Crusafont Pairó
Crusafont Pairó M. (1954). Quelques considérations biologiques sur le Miocène espagnol, Annales de Paléontologie, 40, 97-103.