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La province greco-irano-afghane et la répartition des faunes mammaliennes au Miocène supérieur
... Late Miocene: This is the only sub-epoch with as many as eight reported localities where both subfamilies were found together. All of these fossil sites are located in the Greco-Iranian zoogeographical province (sensu Bonis et al. 1992), herein referred to as the Balkan-Iranian zoogeographical province that includes Southeast Europe, and Anatolia as far as Iran, following Spassov et al. (2018). ...
... Interestingly, the schizotheriine taxon is consistently identified as Ancylotherium pentelicum, which is the only schizotheriine taxon from the Turolian of the Balkan-Iranian province. However, the chalicotheriine material is much more difficult to determine, due to its complicated taxonomic history and fragmentary remains and the existence of up to three different taxa has been suggested (see also Bonis et al. 1992;Anquetin et al. 2007). ...
... In the Eastern Mediterranean specifically, an important trend of cooling and aridification has been observed during the latest Tortonian and earliest Messinian (Mertz-Kraus et al. 2008;Kontakiotis et al. 2019). Furthermore, changes in the sea-level of the Paratethys affected the climate in the wider area and most probably shaped the habitats of the Balkan-Iranian zoogeographical province (Butiseacă et al. 2021;Böhme et al. 2021;Palcu et al. 2021), for which a savannah-like palaeoenvironment has been reconstructed (Abel 1922;Bonis et al. 1992;Böhme et al. 2017;Kaya et al. 2018;Fortelius et al. 2019). Despite the fact that some researchers have argued against this hypothesis, based on the faunal and floral composition (Solounias et al. 1999;Denk et al. 2018), the landscape was marked by the existence of open grasslands with the gradual expansion of C4 plants (Cerling et al. 1993;Tzanova et al. 2015). ...
Chalicotheres are a peculiar group of large herbivorous mammals, closely related to extant tapirs, rhinoceroses, and horses, but with large claws instead of hooves. The family Chalicotheriidae consists of two subfamilies, the Schizotheriinae and the Chalicotheriinae. Herein we present chalicothere remains from the Upper Miocene locality of Pogana 1 in Romania, identifying the schizotheriine Ancylotherium pentelicum and an indeterminate chalicotheriine that were both found in the same stratigraphic layer. Thus, the Pogana 1 locality represents one of the very few confirmed cases of the co-occurrence of the two subfamilies within one fossiliferous horizon in the same fossil site. A detailed review of all localities where the two subfamilies have been reported to co-occur shows that this is a rare phenomenon that is almost exclusively observed in the Turolian of the Balkan-Iranian zoogeographical province. This is probably due to provincial differences in the palaeoenvironment. The data presented here support the hypothesis of a diverse mosaic environment in the Balkan-Iranian province with both closed environments and open habitats that was able to sustain a rich and diverse large mammal fauna.
... De Mecquenem (1924)(1925) identified several giraffid species from the Maragheh collection in the MNHN, Paris, including a braincase of ''Camelopardalis attica'' which is now assigned to Alcicephalus neumayri ( Solounias and Danowitz, 2016a). Maragheh giraffids were later analysed by other paleontologists like Bohlin (1926). Erdbrink (1976aErdbrink ( , 1976bErdbrink ( , 1977Erdbrink ( , 1978) studied the Maragheh collections in the Paleontology Museum of Munich and presented a more complex picture of giraffids that is not clearly valid (Solounias and Danowitz, 2016a). ...
... The lower p4 is molarized. The metapodial bones are elongated (Bohlin, 1926;Geraads, 1974Geraads, , 1978Geraads, , 2009Kostopoulos et al., 1996;Ríos et al., 2016). Fig. 6 is a diagram comparing proportions of PRCI/M352-M353 with B. attica from Pikermi and Nikiti, based on the skull measurements ( Table 1). ...
... Honanotherium is not well Table 1 for skull measurements M1-23. known but it is a more massive animal with shorter metapodials than Bohlinia (Bohlin, 1926). The upper premolars are simple in Honanotherium whereas the styles are curved inward in Bohlinia. ...
... N. Spassov et al. 1974a, b, 1976a, b, 1985, 1989a, b, 1991, 1992, Garevski & Zapfe 1983, Forsten & Garevski 1989, Garevski & Mladenovski 2006, but were mostly published in local journals with limited distribution into the scientific world, some of the articles being even written in native language making all these works of limited use. Some publications of broader diffusion appeared recently (Geraads et al. 2008, Geraads 2009, Garevski & Markov 2011, 2015, Garevski et al. 2012, Radović et al. 2013), but they focused on specific elements of the assemblages, so that the composition, biochronology, and zoogeographical affinities of these late Miocene faunas remain poorly documented compared to other areas of the Balkano-Iranian zoogeographic province (Bonis et al. 1992, Geraads et al. 2003, Spassov et al. 2006, Koufos 2013. ...
... However, their faunal composition was not uniform. Faunal similarities (especially regarding spiral-horned bovids and giraffids) support the existence of a Balkan-Iranian (Greco-Irano-Afghan) zoogeographic province (Bonis et al. 1992), whose westernmost part is now better known, thanks to the field work conducted in the late Miocene of the Republic of Macedonia and our analyses of the resulting collections. The Northern peri-Pontic region shares a number of similarities with this Balkan-Iranian province, especially regarding the hipparions, giraffids, bovids (spiral-horned antelopes, boselaphins, gazelles) (Korotkevich 1988, Krakhmalnaya 1996, and it may be that it was part of this mega-province as well, further research may refine the characteristics of the various sub-provinces. ...
This study represents the first extensive systematic investigation of the Miocene mammalian faunas of the Republic of Macedonia (FYROM), stored in the Macedonian Museum of Natural History, Skopje. They range in age from perhaps the early Miocene to the early Ruscinian, but the bulk of the fossils represent middle Turolian mammals. At least 57 taxa have been identified, from 25 different paleontological sites, mostly from the Vardar and Strumitsa river basins, but also from the Morievo and Delchevo regions. The richest localities are the middle Turolian localities of Karaslari (with 22 species) and Kiro Kuchuk (17 species). The rich fossil material greatly improves our knowledge of the Turolian Hipparion faunas of the Balkan-Iranian zoogeographic paleo-province, whose westernmost part was mostly documented in Greece and Bulgaria. The fauna displays the typical faunal features of the Balkan Pikermian biome, with dominance of hipparions (especially H. brachypus, but our revision does not confirm the presence of Hipparion verae in the Turolian faunas) and bovids such as Gazella, Tragoportax, and spiral-horned antelopes. Other forms usually found in the area, such as Microstonyx erymanthius, Dihoplus pikermiensis, chalicotheres, Choerolophodon pentelici, Mesopithecus pentelicus, or Adcrocuta eximia are also common. Several new forms have been identified among the carnivores, the giraffids and the bovids. The Macedonian material contributes to reconstructing the history of several taxa such as Simocyon, Metailurus, several hipparion species, Propotamochoerus, Bohlinia, Sivatherium. The most noticeable features of these Turolian faunas are: the abundance of spiral-horned antelopes, and rarity of antelopes of the Protoryx-Pachytragus group, as in Bulgaria, the co-existence of chalicotheriins and schizotheriins, the frequency of Dihoplus compared to Ceratotherium, the presence of Chilotherium, which reaches its westernmost longitude, and the presence of Anancus sp. in some localities, considered here as post-Pikermian. © 2018 E. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart, Germany.
... These differences are well expressed in the hipparions of Central and Western Europe. During the Vallesian in the Eastern Mediterranean, the environmental conditions were drier and more open than those of Central and Western Europe (Bonis et al. 1992(Bonis et al. 1999 and consequently the differences between the hipparions from these areas are more expressed. A decrease in the abundance of hipparions is observed during the Pliocene. ...
... Asia Minor was the most continental part of the area between the Balkans and India and perhaps more drier, thus preventing Mesopithecus to live there. In fact, Mesopithecus was present in the two extremes of the Greco-Iranian province (Bonis et al. 1992) near the big mountain chains of the Alps and Himalayans, where the environment was probably less dry. ...
... The geographic area that presently incorporates the southern Balkans, Turkey, and part of the south peri-Pontic region formed during the Late Miocene western domain of the so-called Greco-Iranian biogeographic province (Bonis de et al., 1992). The fossils of Hayranlı in this study were from the Derindere region of the İncesu formation, from the Sivas plain, belonging to the Greco-Iranian biogeographic province. ...
... According to their biochronological relationships, the Sivas carnivores expanded mainly between European MN11 and 12, or 8.7-6.6 million years ago. The geographic area that presently incorporates the southern Balkans, Turkey, and part of the southern peri-Pontic region formed the western domain of the so-called Greco-Iranian biogeographic province during the Late Miocene (Bonis de et al., 1992). Kostopoulos (2009) emphasized that the early Turolian large mammal association of southeastern Europe, although it followed general trends, seemed to absorb most of the environmental vibrations that provided more vigorous reactions in the eastern sector (i.e. ...
The locality of Hayranlı-Sivas is situated in the central Anatolian plateau (Sivas, Turkey) and includes many fossil remains, including carnivores collected from 2 different spots. The aim of this study is to contribute to knowledge of carnivore evolution in Turkey based on the fossil specimens in Hayranlı-Sivas. The studied collection indicates the presence of the following taxa: Hyaenictitherium wongii, Hyaenictitherium intuberculatum, Lycyaena dubia, and Machairodus giganteus. L. dubia is the first record from Anatolia. The material of each taxon was described and determined by comparison with other materials from various Eurasian localities. During the Early or Middle Turolian 9–7 Ma. (MN11–12), shrubland and open savanna grassland ecosystem habitats might have contributed to rich faunal diversity in the Hayranlı location. Moreover, carnivores of the area, represented by 4 taxa biochronologically, were adapted to this ecology during the evolutionary processes. M. giganteus in closed ecosystem locality HAY-91 and H. wongii, H. intuberculatum, and L. dubia in open ecosystem locality HAY-2 were probably the most dominant carnivores of the survey area.
... Most probably they originate from Turolian migrants from the Balkans or Asia Minor (Begun et al. 2003), which were preadapted to the conditions of relatively sclerophyllous forest vegetation. Th is would support the idea of Solounias et al. (1999) that the origin of the modern African fauna of the sclerophyllous savannahs can be found in the "Pikermian biome" of the so-called Balkano-Iranian (Greco-Iranian) (Bonis et al. 1979(Bonis et al. , 1992b or subparatethyan (Bernor 1983) zoogeographic province. Th e aridifi cation at the end of the middle Turolian could be the main cause of the hominoid extinction from the Balkano-Anatolian region and their survival under less severe conditions in Africa. ...
... Rare reddish horizons in the Strumyani deposits provide additional indications about this aridifi cation. Th is is the time of the climax of the so-called "Pikermian biome" sensuSolounias et al. (1999) on the territory of the Greco-Irano-Afghan (sensuBonis et al. 1992b) (i.e. the Balkano-Iranian) faunal paleo-province and the time of maximal dispersal of a fauna similar in appearance to that of the recent African savannah. Th e occurrence of many mammalian fossil localities in fl uviatile context within the caldera of the Kojuh paleo-volcano (such as Kromidovo-1, Kromidovo-2, Vinogradi and Marino Pole) indicates the complete drying up of the previous lake in this area, although fragmented swamps still persist on a large part of its territory. ...
The stratigraphy of the Neogene deposits along the Middle Struma River (SW Bulgaria) is revised. Five genetic lithocomplexes are recognized, replacing the numerous lithostratigraphic formations currently used. The basic concept is that the upper Miocene alluvial-proluvial deposits along the Middle Struma River Valley are a product of "braided rivers" transporting important volumes of loose rocks, characterized by rapid and irregular accumulation. This accumulation occurred in conditions of increasing aridification. Uprising of the Rila and Pirin Mountains occurred later. Unlike northern Greece, where Vallesian landscapes were relatively open, humid habitats and forest vegetation seem to be well represented in the late Vallesian of the middle Struma region, but herbaceous and shrub communities already had a significant role. The Mesta River might have flowed into the Middle Struma, south of today's Petrich tectonic basin. The Turolian landscape can be reconstructed thanks to the rich Turolian mammalian faunas from the numerous (about 40) localities along the Middle Struma. Dominant taxa are Palaeoreas lindermayeri, Hipparion (H. brachypus, H. gr. mediterraneum-moldavicum, and H. cf. macedonicum), Gazella, Tratgoportax, giraffes, Their likely ecological requirements show that the "Pikermian biome" (sensu Solounias et al. 1999) was dominated by open woodlands resembling park type forest (rather than by shrubby vegetation). By the end of the middle and the beginning of the late Turolian, time of accumulation of Piperitsa Genetic Lithocomplex, characterized by reddish terrigenous-sandy deposits, open landscapes probably prevailed, with spots of sclerophyllous woodlands. This is probably the time of the first occurrence of the genus Anancus in the middle Struma, Bulgaria, and Europe. © Publications Scientifiques du Muséum national d'Histoire naturelle.
... These differences are well expressed in the hipparions of Central and Western Europe. During the Vallesian in the Eastern Mediterranean, the environmental conditions were drier and more open than those of Central and Western Europe (Bonis et al. 1992(Bonis et al. 1999 and consequently the differences between the hipparions from these areas are more expressed. A decrease in the abundance of hipparions is observed during the Pliocene. ...
... Asia Minor was the most continental part of the area between the Balkans and India and perhaps more drier, thus preventing Mesopithecus to live there. In fact, Mesopithecus was present in the two extremes of the Greco-Iranian province (Bonis et al. 1992) near the big mountain chains of the Alps and Himalayans, where the environment was probably less dry. ...
A biostratigraphic division of the Neogene in the Eastern Mediterranean is highly necessary. Data from Late Miocene mammalian faunas and absolute dating were collected and used for a first bio-stratigraphic division. Some important mammalian taxa have been selected and their paleobioge-ographic distribution is given. The dispersal events of these taxa are used for the division of the Late Miocene. A preliminary biozonation of the Late Miocene is proposed based on mammalian faunas. However, more data from various countries are necessary in order to obtain a quite reliable biostratigraphy.
... At the beginning of the 1980s little was known about the Late Miocene carnivores of the Greco-Iranian Province (Balkans, Turkey, Iran and Afghanistan), (Bonis et al., 1992a). Most of the available data were based on the old collections from Pikermi, Samos and Maragheh. ...
... At the beginning of the 1980s little was known about the Late Miocene carnivores of the Greco-Iranian Province (Balkans, Turkey, Iran and Afghanistan), ( Bonis et al., 1992a). Most of the available data were based on the old collections from Pikermi, Samos and Maragheh. ...
The Late Miocene carnivore assemblages of the Greco-Iranian Province (Balkans, Turkey, Iran and Afghanistan) are analyzed by different methods to determine their relationships and palaeoecology. The carnivores are separated into five assemblages according to their MN age (European Neogene Mammal Zones). Faunal composition, three-dimensional guild structure diagrams and multivariate analysis are used for their study and comparison. The Late Miocene carnivore assemblages of the studied area are characterized by the dominance of hyaenids, while percrocutids and felids are more abundant in the Vallesian and Turolian, respectively. The predators of the Greco-Iranian Province were generally living in an open environment, as indicated by the analysis and comparison of their guild structure with modern assemblages. The guild structure diagrams of all carnivore assemblages resemble to that of the modern Serengeti one, corresponding to an open environment. Similar results about their environments were obtained from their multivariate analysis and comparison with modern assemblages from known environments. The Late Miocene carnivore assemblages match together in a group which can be clearly correlated to the modern assemblages from open habitats. There are, however, some small differences between the Vallesian (early Late Miocene) and Turolian (late Late Miocene) palaeoenvironmental conditions. The Vallesian palaeoenvironment was relatively wetter and more closed than that of the Turolian. The Turolian is characterized by an increase of the open character until the late Turolian, when there are indications for wetter and more closed conditions, as indicated by the guild structure of the MN 13 assemblage.
... Most probably they originate from Turolian migrants from the Balkans or Asia Minor (Begun et al. 2003), which were preadapted to the conditions of relatively sclerophyllous forest vegetation. Th is would support the idea of Solounias et al. (1999) that the origin of the modern African fauna of the sclerophyllous savannahs can be found in the "Pikermian biome" of the so-called Balkano-Iranian (Greco-Iranian) (Bonis et al. 1979(Bonis et al. , 1992b or subparatethyan (Bernor 1983) zoogeographic province. Th e aridifi cation at the end of the middle Turolian could be the main cause of the hominoid extinction from the Balkano-Anatolian region and their survival under less severe conditions in Africa. ...
... Rare reddish horizons in the Strumyani deposits provide additional indications about this aridifi cation. Th is is the time of the climax of the so-called "Pikermian biome" sensuSolounias et al. (1999) on the territory of the Greco-Irano-Afghan (sensuBonis et al. 1992b) (i.e. the Balkano-Iranian) faunal paleo-province and the time of maximal dispersal of a fauna similar in appearance to that of the recent African savannah. Th e occurrence of many mammalian fossil localities in fl uviatile context within the caldera of the Kojuh paleo-volcano (such as Kromidovo-1, Kromidovo-2, Vinogradi and Marino Pole) indicates the complete drying up of the previous lake in this area, although fragmented swamps still persist on a large part of its territory. ...
Th e stratigraphy of the Neogene deposits along the Middle Struma River (SW Bulgaria) is revised. Five genetic lithocomplexes are recognized, replacing the numerous lithostratigraphic formations currently used. Th e basic concept is that the upper Miocene alluvial-proluvial deposits along the Middle Struma River Valley are a product of "braided rivers" transporting important volumes of loose rocks, characterized by rapid and irregular accumulation. Th is accumu- lation occurred in conditions of increasing aridifi cation. Uprising of the Rila and Pirin Mountains occurred later. Unlike northern Greece, where Vallesian landscapes were relatively open, humid habitats and forest vegetation seem to be well represented in the late Vallesian of the middle Struma region, but her- baceous and shrub communities already had a signifi cant role. Th e Mesta River might have fl owed into the Middle Struma, south of today's Petrich tectonic basin. Th e Turolian landscape can be reconstructed thanks to the rich Turolian mammalian faunas from the numerous (about 40) localities along the Middle
... The Late Miocene of Eurasia is an especially interesting time period for fossil rhinos, with several species being able to coexist in the same locality (Antoine, in press;Antoine & Saraç, 2005;Deng, 2006b;Fortelius et al., 2003;Heissig, 1999;Pandolfi, 2016). The Balkan-Iranian province (Bonis et al., 1992;Kampouridis et al., 2023;Spassov et al., 2019) is one of the best-studied regions with world-renowned Upper Miocene fossil localities such as Pikermi (Gaudry, 1862;Roussiakis et al., 2019;Wagner, 1848) and Samos (Kostopoulos et al., 2003;Koufos et al., 2011;Osborn, 1899;Solounias et al., 2010) in Greece, Hadjidimovo (Hristova et al., 2003;Spassov, 2002;Spassov & Geraads, 2004) in Bulgaria, Karaslari (Garevski, 1974;Schlosser, 1921;Spassov et al., 2018) in North Macedonia, and Maragheh (Ataabadi et al., 2013;Bernor, 1986;Hullot et al., 2022;Mecquenem, 1908;Pohlig, 1886) in Iran. Despite the abundance of material that has been excavated for over a century, many issues continue to exist regarding the rhinos. ...
Extant rhinoceroses are represented only by five species and are characterized by the presence of a nasal horn. In the past, they were much more diverse, with one of the best-known groups being the aceratheriines, i.e., hornless rhinoceroses. Chilotheres are a group of hornless rhinos that inhabited Eurasia during the Late Miocene. Their westernmost geographic range reached Eastern Europe, where overall eight species have been erected. Four of these were described based on material from the Upper Miocene of Samos Island (Greece), two of which are not considered valid anymore. Unfortunately, the type skulls of all four species are lost and there are several issues concerning their taxonomy. Therefore, we herein designate two skulls housed in historical collections from Samos as neotypes for the first two species, Chilotherium schlosseri (Weber, 1905) and Eochilotherium samium (Weber, 1905), and provide detailed comparisons for the separation of the species from each other and from any other chilotheres. Our results prove that the two species are valid and justify their separation on a generic level. Chilotherium schlosseri seems to be more closely affiliated with the other European Chilotherium species, whereas E. samium is more similar to the Chinese 'Chilotherium' wimani and 'Chilotherium' primigenium, based on their more plesiomorphic characters.
... Samos was part of Asia Minor during the Late Miocene and its faunal assemblages show more similarities with the western Asian assemblages compared with the southern Balkans ones (Kostopoulos, 2009;Koufos et al., 2009). In terms of proboscideans, the collective Samos fauna includes all typical Turolian members of the Greco-Iranian-Afghan (sensu Bonis et al., 1992; Balkano-Iranian or Sub-Paratethyan) paleobiogeographic province: Deinotherium proavum, "Mammut" sp. ("M." obliquelophus?), Choerolophodon pentelici, and Konobelodon atticus (Konidaris & Koufos, 2019). ...
Mammutidae comprise a proboscidean family that originated in Africa during the late Oligocene, dispersed across the Holarctic during the Miocene, and survived in North America until the end of the Pleistocene. Despite their long evolutionary history and wide geographic distribution mammutids are particularly scarce in the Miocene of Eurasia. Here, we present a new mammutid specimen (an upper deciduous premolar) from the Upper Miocene locality of Sazak in southwestern Turkey. Morphological and metric traits of the tooth, in particular the well-expressed zygodonty, are distinct from the more basal Zygolophodon and permit its assignment to the more derived “Mammut.” Due to the absence of more diagnostic specimens, a specific attribution is not possible; however, considering the Turolian age of the associated fauna an attribution to the Late Miocene representative of the genus, “Mammut” obliquelophus, is possible. Turolian mammutids are rare in the fossil record and therefore our knowledge remains only fragmentary. Despite the existence of a single specimen, the presence of this genus in Sazak corresponds to its first report in the Upper Miocene of Turkey, as well as the first one in western Asia. The presence of “Mammut” in the Upper Miocene of China was recently confirmed, and therefore the record of “Mammut” at Sazak, i.e., at the western margin of Asia, not only adds to the scanty record of the genus in the Upper Miocene of Eurasia but also provides another line of evidence of the paleozoogeographic link enabling Europe–East Asia proboscidean interchanges during the Late Miocene.
... They share many morphological features related to their adaptation to cursorial life. Struthio is the only genus of this group present in the late Miocene of the sub-Paratethyan bioprovince (sensu Bernor 1983) or Greco-Iranian (Balkan-Iranian) province (sensu de Bonis et al. 1992). Nonetheless, Struthio was not the only cursorial bird that existed during the late Miocene in the sub-Paratethyan (Balkan-Iranian) bioprovince. ...
The present study describes an almost complete cervical region of the fossil ostrich Struthio karatheodoris from the Turolian locality of Kerassia (Euboea, Greece). The material comes from two distinct fossiliferous horizons and consists of twelve cervical vertebrae, ten of which belong to the same individual. These specimens are the first remains of a large flightless bird from Kerassia and represent some of the very few findings of the genus Struthio in the sub-Paratethyan (Balkan-Iranian) bioprovince from the late Miocene. The morphology of the cervical vertebrae implies that the neck of S. karatheodoris had similar biomechanical properties to that of extant ostriches, pointing to similar ecological adaptations, relating to food procurement. However, it probably had a more flexible and stronger neck, which might indicate a somewhat different ecology. Furthermore, considerable intraspecific size and morphological variation of the cervical vertebrae of S. karatheodoris is observed. This signifies that S. karatheodoris was not necessarily larger than S. camelus as previously suspected, but their size ranges in fact overlap significantly.
... 9-8 Ma) (Sen et al. 1997). Geographically, Molayan and Taghar are situated close to the border between two paleobiogeographical provinces, the Greco-Iranian and the Siwalik provinces (de Bonis et al. 1992). However, the mammalian faunas from these two Afghan localities are firmly Greco-Iranian in composition, with hardly a species in common with the Potwar Plateau only some 300 km to the southeast (Brunet et al. 1984). ...
... 9-8 Ma) (Sen et al. 1997). Geographically, Molayan and Taghar are situated close to the border between two paleobiogeographical provinces, the Greco-Iranian and the Siwalik provinces (de Bonis et al. 1992). However, the mammalian faunas from these two Afghan localities are firmly Greco-Iranian in composition, with hardly a species in common with the Potwar Plateau only some 300 km to the southeast (Brunet et al. 1984). ...
Modern day South and Southeast Asia falls almost entirely within a single biogeographic region, the Indomalayan Realm. Here, we review the Cenozoic geological and environmental history of the Indomalayan Realm, and the evolutionary history of the large herbivorous mammals that have inhabited it. For the most part bounded by major physical features, the cohesiveness of the Indomalayan Realm in its mammalian faunas can be recognized as far back as the middle Miocene or even earlier. Many of the extant large herbivores of the Indomalayan Realm have a diverse fossil record in this part of the world, though a few, such as cervids, are relative newcomers. Many extant clades that are not currently present in the Indomalayan Realm had records in the region up to the Pleistocene, including giraffids, hippopotamids, and reduncin antelopes. The island archipelago of Southeast Asia in particular witnessed radiative speciation of numerous clades including proboscideans
and ruminants, at least through the climatic cycles of the Pleistocene, if not earlier. If there is a single common thread to the evolutionary history of Indomalaya’s large herbivores, it may be the loss of taxonomic diversity, with much greater taxonomic representation recorded at numerous times in the past in almost all large herbivore clades. Today, diversity loss continues at the hands of anthropogenic, rather than natural, environmental causes, which threaten to violently curtail millions of years of evolutionary heritage.
... There is another reference about its presence in the locality Novoukrainka, Ukraine (Pidoplichko, 1959ex Coombs, 1989, where Pidoplichko described an articulating Mc II , duplex and large distal phalanx, coming from a hipparion fauna, which seems to be similar to A. pentelicum. The species is quite rare in Western Europe, but it is well distributed in the Balkan Peninsula and Turkey; it is also reported from Maragheh, Iran (de Mecquenem, 1924-25) and Molayan, Afghanistan (Sen, 1998), the Greco-Iranian Province as it was defined by Bonis et al. (1992). Considering the similarities of the Chinese material with A. pentelicum, it is quite possible that this material belongs to this species, expanding its geographic distribution to the whole Asia. ...
... The oldest evidence of A. pentelicum is known from the Vallesian locality of Pentalophos in Greece, but the material is scanty (Bonis et al., 1999). During the Turolian, the geographic range of A. pentelicum covers the entire Subparatethyan (Bernor, 1983(Bernor, , 1984 or Greco-Iranian (Bonis et al., 1992a(Bonis et al., , 1992b zoogeographic province (Fig. 1). It has been firmly documented in the localities of Pikermi (Wagner, 1857;Hensel, 1862;Gaudry, 1863;Thenius, 1953;roussiakis & Theodorou, 2001), Samos (Major, 1894;Schaub, 1943), Halmyropotamos (Melentis, 1969a(Melentis, , 1969b, Kerassia (Theodorou et al., 2004), and Thermopigi (Geraads et al., 2007) in Greece; Veles in Fyr of Macedonia (Schlosser, 1921;Garevski, 1974;Garevski & Zapfe, 1983); Gorna Sushitsa, Kalimantsi, Hadjidimovo, and Strumyani-1 in Bulgaria (Bakalov, 1955;Bakalov & Nikolov, 1962;Geraads et al., 2001Geraads et al., , 2006; Gülpinar (Kaya, 1986), Kemiklitepe (Kaya, 1988;Sen, 1994), Salihpaşalar and Pinaryaka (Saraç et al., 2002), Karaburun (Kaya et al., 2005), Konya-Kızılören and Akkaşdaği (Saraç & Sen, 2005) in Turkey; and Maragheh in Iran (Mecquenem, 1924;Geraads et al., 2006). ...
... tinent by diverse taxa, including catarrhines and thryonomyids. Faunal similarity appears to be marginally greater among low-latitude areas of Africa and southern Asia than with more temperate regions to the north, which illustrates the distinction of the temperate Greco-Irano-Afghan faunas (de Bonis et al. 1992) and (through time) the Pikermian chronofauna (Eronen et al. 2009). ...
This chapter analyzes the fossil record of South Asia's Neogene small land mammals in the context of its paleobiogeography. Within the Indian Subcontinent, high faunal similarity can be observed among local small mammal assemblages distributed on a scale of 1000 kilometers, and these are distinct from assemblages to the west and northwest and northeast, beyond the subcontinent. Eastward into Thailand, Myanmar (Burma), and Yunnan, China, faunal similarity with the Indian Subcontinent is apparent but weaker than within the subcontinent. This pattern mimics the distribution of the present-day Oriental biogeographic province. This chapter examines the extent to which this paleozoogeographic pattern can be defined, as well as faunal elements that do not follow the rules. Evidence from Yunnan, Thailand, and Myanmar shows that many genera of small mammals are shared throughout southern Asia.
... The Turolian fauna of the Greco-Iranian Province (Bonis et al., 1992b) was considered as homogeneous for a long time (Bernor et al., 1996;Solounias et al., 1999Solounias et al., , 2010. Recently the study and analysis of the Turolian mammal assemblages from both sides of the Aegean Sea indicated that several tectonic events and palaeoclimatic changes affected the area, causing the isolation of the Southern Balkans (western Aegean domain, WAD) from Samos and Asia Minor (eastern Aegean domain, EAD) during the Turolian. ...
... The estimated divergence times imply speciation of A. witherbyi at the Miocene/Pliocene transition (about 6 Ma). The late Miocene was characterized by extended savannah, called the Greco-Iranian Province, from the Balkan Peninsula to Iran and Afghanistan (De Bonis et al. 1993). During the late Miocene, seasonal rainfall led to the establishment of grasslands with small trees and replacement of the woodland by more open habitats (Koufos and Konidaris 2011). ...
Wood mice of the genus Apodemus are widely distributed in Eurasia, with the Eastern Mediterranean being considered as a hotspot. Indeed, numerous species have been documented in Iran, including A. witherbyi, A. hyrcanicus, A. uralensis, A. avicennicus, A. hermonensis, and A. arianus. In this study, 129 specimens were collected from different Iranian localities and two specimens from Afghanistan. The animals were identified taxonomically and their phylogenetic relationships were investigated using cytochrome b mitochondrial DNA sequences. Five species of the genus Apodemus were identified in Iran, including A. hyrcanicus, A. witherbyi, A. cf. ponticus, A. uralensis, and A. mystacinus, beside, A. pallipes from Afghanistan. This study found no evidence of A. flavicollis or A. sylvaticus in Iran, despite their occurrence in Turkey, shedding doubt on the status of A. flavicollis in Iran, Asia Minor, and the Levant. Phylogenetic analyses imply that A. witherbyi has priority over A. avicennicus, A. hermonensis, and A. iconicus. Estimation of the divergence time for these taxa suggests a separation at around 7.2 Ma for the subgenera Karstomys (including A. mystacinus and A. epimelas) and Sylvaemus (including A. flavicollis, A. sylvaticus, A. uralensis, A. pallipes, A. hyrcanicus, A. witherbyi, and A. cf. ponticus). Within the subgenus Karstomys, the divergence times for A. mystacinus andA.epimelas were between 3.0 and 6.1 Ma, and divergence times within the subgenus Sylvaemus were between 5.2 and 6.9 Ma forA.witherbyi and other species in this subgenus. It is postulated that vicariance events including the uplifting of the Zagros Mountains and Anatolian Plateau in the middle Miocene and climate oscillations during the Messinian Salinity Crisis besides formation of the Hyrcanian tertiary forests during the Neogene probably played substantial roles in the radiation and distribution of the genus Apodemus in the Eastern Mediterranean.
... Irano-Anatolian and Caucasus hotspots are two of 35 regions proposed for conservation priorities (Myers et al., 2000;Mittermeier et al., 2012). Additionally, the northwest of Iran was a part of paleo-corridor between Iran and Anatolia where provided exchange of mammals between Europe and Asia (Wessel, 1955;De Bonis et al., 1993). Particularly, the region represents a high diversity of terrestrial mammals which may be attributed to its diverse ecological conditions. ...
The northwest Iran comprises parts of two major biodiversity hotspots; Irano-Anatoli and Caucasus. It is a mountainous transition realm between Caucasus forest in the north, Pontic forest of Turkey in the west and central deserts of Iranian Plateau. This study was designed to determine rodent diversity in northwest Iran. Moreover, corridor and barrier features of the region were investigated, as well. The samplings were done in different localities of northwestern Iran. In addition, all specimens of the Zoology Museum of Ferdowsi University of Mashhad (ZMFUM) attributed to the region were denoted. The study shows that the specimens belong to 18 nominal forms attributing to 5 families: Muridae (Apodemus witherbyi, Mus musculus domesticus, Mus macedonicus, Meriones persicus rossicus, M. libycus erythrourus, M. vinogradovi and M. tristrami bogdanovi), Gliridae (Dryomys nitedula pictus), Cricetidae (Microtus socialis, M. obscurus, M. qazvinensis, Chionomys nivalis trialeticus, Arvicola amphibious persicus, Ellobius lutescens lutescens, Cricetulus migratorius pulcher and Mesocricetus brandti brandti), Dipodidae (Allactaga williamsi schmidti) and Calomyscidae (Calomyscus urartensis).
... Th ey therefore provide further support for the existence of faunal and environmental similarities across a larger geographical region from the Balkans to Iran and Afghanistan, the so-called Southeast European-Southwest Asian "superprovince" sensu Bernor et al. (1996) or the Greco-Iranian (i.e. Balkan-Iranian) province sensu Bonis et al. (1992), during that period. ...
We describe two new fossils, the distal end of a right tarsometatarsus and a proximal pedal phalanx of the left third toe, from two sites in southwestern Bulgaria: Kalimantsi (middle Turolian) and Hadzhidimovo-2 (MN 11/12 boundary). These specimens are compared to Neogene-Quaternary ostriches, and are referred to Struthio cf. karatheodoris. A general overview of Neogene-Quaternary ostrich specimens, a taxonomic discussion of late Miocene Eurasian struthionid taxa, and the ecological and zoogeographic implications of the new specimens are presented.
... Previous studies on late Miocene faunal assemblages point out strong environmental differences between western, central, and eastern Europe [Bernor, 1984;Bonis et al., 1992aBonis et al., , 1992bBonis et al., , 1992cFortelius et al., 1996]. Ecological indicators from hypsodonty index to diversity analysis to body weight distribution of large herbivorous mammals indicate more open landscapes like wooded savannah or flood plain grasslands during the Vallesian (MN 9/10) from eastern Europe to Anatolia [Bonis et al., 1992a;Fortelius et al., 1996Fortelius et al., , 2002. ...
This study deals with the paleoenvironmental changes in northern Greece during the late Vallesian and the early Turolian,
before and after the last occurrence of the hominoid Ouranopithecus macedoniensis. Dental microwear of fossil bovids yields information on paleodiet; thus, inferences can be drawn about floristic composition
and paleoenvironmental changes. The microwear pattern of the fossil species is compared with that of extant species from the
database “Ungulates” (20 extant species and 471 wild-shot specimens) in a multivariate analysis. The bovids of the late Vallesian
“Ravin de la Pluie” locality show a dental microwear pattern similar to that of the extant grazers. This attests to the presence
of open landscapes with an important grassy herbaceous layer in northern Greece during the late Vallesian. The bovids from
the early Turolian also grazed. Nevertheless, Tragoportax rugosifrons, which constitutes the largest sample from the Turolian “Ravin des Zouaves 5” locality, was a mixed feeder. These dietary
adaptations indicate an environment of bushy and/or wooded areas with a grassy herbaceous layer. The abundance of the mixed
feeders in the ungulate assemblage of this latter locality also points out strong seasonal fluctuations in food availabilities.
... Some premolars have large anterior lobe (paraconid and parastylid) while a few have small anterior lobe with closed lingual side. The variation in premolars of Giraffokeryx recorded from Candir and Pasalar (Turkey), and the Siwaliks, have also been observed by the earlier researchers (Pilgrim, 1911;de Bonis et al., 1992;Geraads and Aslan, 2003). It may be suggested that more than one species was present in the Pakistani Lower Siwaliks as noted by Geraads and Aslan (2003). ...
New mammalian material excluding proboscideans described from four main fossil sites of the
Chinji Formation, northern Pakistan, allow identifying Miotragocerus gluten, Tragoportax cf. punjabicus,
Elachistoceras sp., Helicoportax sp., Boselaphini sp. indet., Gazella sp., Giraffokeryx punjabiensis, Giraffa priscilla,
Dorcatherium minus, Microbunodon silistrensis, Merycopotamus nanus, Listriodon pentapotamiae, Conohyus
sindiensis, Gaindatherium browni and Hespanotherium matritense. The tooth positions of all the fifteen species are
documented. The findings enlarge our knowledge on the anatomic features of the described species. Quantitatively,
the bovid taxa are the most predominant and provide existing evidence of small (Elachistoceras) and medium
(Miotragocerus, Tragoportax, Helicoportax, Gazella) size bovids with other Middle Miocene mammalian taxa. The
composition of the fauna suggests a Middle Miocene age. The palaeoclimate of this age (14.2 and 11.2 Ma) can be
interpreted warm and humid having extensive forest component and developed grassy sub areas.
... The Late Miocene rhino assemblages of Greece are in conformity with the ones from Southeastern Europe and the Eastern Mediterranean, especially with the ones from Anatolia. From a biogeographical aspect, these results correspond to the Eastern regional block of Fortelius et al. (1996) or, in a closer approach, to the Greco-Iranian or sub-Paratethyan mammalian province of Bernor (1984) and De Bonis et al. (1993, 1999. By the end of the Turolian, all these species became extinct in Greece, unable to survive the paleoenvironmental changes and the faunal turnover at the Miocene-Pliocene boundary. ...
... The oldest evidence of A. pentelicum is known from the Vallesian locality of Pentalophos in Greece, but the material is scanty (Bonis et al., 1999). During the Turolian, the geographic range of A. pentelicum covers the entire Subparatethyan (Bernor, 1983(Bernor, , 1984 or Greco-Iranian (Bonis et al., 1992a(Bonis et al., , 1992b zoogeographic province (Fig. 1). It has been firmly documented in the localities of Pikermi (Wagner, 1857;Hensel, 1862;Gaudry, 1863;Thenius, 1953;roussiakis & Theodorou, 2001), Samos (Major, 1894;Schaub, 1943), Halmyropotamos (Melentis, 1969a(Melentis, , 1969b, Kerassia (Theodorou et al., 2004), and Thermopigi (Geraads et al., 2007) in Greece; Veles in Fyr of Macedonia (Schlosser, 1921;Garevski, 1974;Garevski & Zapfe, 1983); Gorna Sushitsa, Kalimantsi, Hadjidimovo, and Strumyani-1 in Bulgaria (Bakalov, 1955;Bakalov & Nikolov, 1962;Geraads et al., 2001Geraads et al., , 2006; Gülpinar (Kaya, 1986), Kemiklitepe (Kaya, 1988;Sen, 1994), Salihpaşalar and Pinaryaka (Saraç et al., 2002), Karaburun (Kaya et al., 2005), Konya-Kızılören and Akkaşdaği (Saraç & Sen, 2005) in Turkey; and Maragheh in Iran (Mecquenem, 1924;Geraads et al., 2006). ...
In the present study, we describe several postcranial elements of Ancylotherium pentelicum (Chalicotheriidae, Schizotheriinae) from the island of Samos (Greece), some of which are recorded for the first time from this site. The material is compared to the known hypodigm of the species, as well as with other chalicotheres. The known Miocene record of the two chalicothere subfamilies, Chalicotheriinae and Schizotheriinae, in the Eastern Mediterranean and adjacent regions is briefly reviewed and evaluated. The potential paleoecological preferences of both subfamilies are briefly discussed. The presence of the schizotheriine A. pentelicum and the absence of any chalicotheriine species in the Turolian faunal assemblages of Samos and Anatolia indicate the expansion of more open habitats in these localities compared to continental Greece, Fyr of Macedonia and Bulgaria.
... Chalicotheres are well known in the Greco-Iranian Palaeoprovince (GRIP), a region extended from the Balkan Peninsula to Afghanistan (Bonis et al., 1992a), found in several Miocene localities (Garevski, 1974;Garevski and Zapfe, 1983;Bernor et al., 1996;Sen, 1998;Koufos, 2006a;Geraads et al., 2006Geraads et al., , 2007Saraç, 2003;Saraç and Sen, 2005). The presence of A. pentelicum in the late Miocene locality of Akkas¸da˘Akkas¸da˘ gı is questionable. ...
Chalicotheriids are rare in the late Miocene mammal localities of Axios Valley, Macedonia (Greece). The
new campaign of excavations, since 1972, has provided some specimens, which are studied in this article.
They are coming from two different localities. The late early Vallesian locality of Pentalophos 1 (PNT) has
provided a skull and a mandible of an Ancylotherium. The morphological characters of the PNT material as
the small size, the long snout, the shallow mandibular corpus, the strong cingulum in the teeth, the short tooth
rows and the short M3/m3 indicate that it differs from the known Turolian species A. pentelicum and allow
the erection of a new species, named Ancylotherium hellenicum n. sp., which can be used as a biostratigraphic
marker of the Vallesian. The middle Turolian locality Prochoma 1 (PXM) has provided only one M3, which is
determined to the chalicotheriine Anisodon macedonicus. This species was earlier described from the middle
Turolian locality Vathylakkos 3 (VAT) and the late Turolian one of Dytiko 3 (DKO) of Axios Valley. The
biogeography and biostratigraphy of the late Miocene chalicotheres of the Greco-Iranian Palaeoprovince
(GRIP), as well as their palaeoecology are also discussed. The common chalicothere of GRIP is A. pentelicum,
expanded from the Balkans to Afganistan and ranging stratigraphically from the early to the late Turolian.
... Molayan is one of the richest mammal localities in Afghanistan (Sen, 1998). The occurrence of many common species and genera at Molayan, where more than 4000 specimens of large mammals have been collected, and in several middle Turolian localities of Iran, Turkey and Greece, has led Bonis et al. (1992) to de¢ne a Greco-Irano-Afghan (GIA) biogeographic province during the late Miocene. The fossil assemblage from Molayan is dominated by ruminants, among which the bovid Tragoportax (tribe Boselaphini) is very common. ...
The rich mid-Turolian site of Molayan (Afghanistan) has yielded more than 100 mandibles from the bovid Tragoportax afghanicus. In this study we document different aspects of the paleobiology and paleoecology of this fossil bovid by examining patterns of inter- and intra-tooth carbon and oxygen isotopic analyses in M1 to M3 molars from five specimens. Accurate paleoecological and paleoclimatic inferences based on carbon and oxygen isotope analyses in fossil enamel are directly dependent upon the way in which teeth are sampled as well as taphonomic processes which are responsible for fossil accumulation. Because this sampling methodology provides a unique opportunity to detect seasonal changes in diet and environment during tooth formation, it also offers the possibility of testing between catastrophic and progressive deposition at this site. Preservation of δ13C and δ18O values is shown to be good and permits inter-individual comparisons of isotopic trends to be made. Timing of molar crown formation and enamel growth rate were deduced from the pattern of intra-tooth carbon and oxygen isotope variation. The pattern of δ18O variation found in the first and second molars is similar in all the individuals, which suggests that a single birth season occurred probably before the hot/dry season. Large differences in average δ18O values (>2.5‰) are found within a single cohort and indicate that fossil accumulation was progressive rather than catastrophic. The hypothesis that T. afghanicus evolved in an open environment is supported by high δ13C values in a C3 context. Increasing temperature and/or aridity eastward along the Greco-Irano-Afghan biogeographic province is strongly suggested by increasing δ18O values between Greece and Afghanistan.
... The Turkish Neogene sedimentary and fossil record is very rich and offers the opportunity for high-resolution sampling and reconstruction of Miocene paleobiotas and paleoenvironments. Lying in what has been termed the Sub-Paratethyan (Bernor, 1983) or Greco-Iranian-Afghan (de Bonis et al., 1992b) paleobiological faunal province, Turkey and the Balkan region's ancient biotas were relatively continuous with those to the east, all the way to Kabul, Afghanistan, and possibly China. The Neogene was witness to the development, evolution, and ultimate demise of such Eurasian faunal provinces against a backdrop of changing environments and climates, including gradually decreasing global temperatures since the mid-Miocene (Zachos et al., 2001). ...
Craniodental remains of fossil bovids from the late Miocene İncesu Formation, from sites near the city of Sivas, Turkey, are described. The bovid remains represent at least five species: Gazella cf. G. capricornis, Prostrepsiceros houtumschindleri syridisi, cf. Protoryx sp., Tethytragus cf. T. koehlerae, and Tragoportax cf. T. amalthea. The Sivas fossil bovid assemblage is fairly typical for the Greco-Iranian-Afghan paleobiological province, and compares well with the classic Turolian sites of Samos, Pikermi, and Maragheh. Biochronological correlations using these fossil bovids suggest the Sivas fossil assemblage is MN11 or early MN12 in age, or somewhere between 9–7 Ma. The presence of Tethytragus at Sivas represents only the second occurrence of this typically mid-Miocene (MN5–8) taxon from the late Miocene. Paleoecological attributes of the Sivas fossil bovids suggest local paleoenvironments at Sivas comprised shrubland to woodland biomes, perhaps devoid of expansive grasslands or dense forests. The presence/absence and relative abundances of bovid taxa within and among different Greek and Anatolian late Miocene fossil sites is compared and contrasted by way of correspondence analysis. Sivas plots among a number of sites all characterized by high proportions of Gazella, Tragoportax, and Protoryx/Pachytragus, and these in turn are readily distinguished from sites in which Palaeoreas/Majoreas, Protragelaphus, Oioceros, and Miotragocerus are more common. It is suggested that Sivas and similar sites (e.g. Sinap, Akkas¸daği) may have sampled drier, more open habitats than those with strongly differing faunal compositions (e.g. Nikiti-1, Çorakyerler).
... 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). These data are further supported by Koufos et al. (2009a: figs. ...
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 (βΌ1200 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 β€four stages-of-evolutionβ€ 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.
... Th ey therefore provide further support for the existence of faunal and environmental similarities across a larger geographical region from the Balkans to Iran and Afghanistan, the so-called Southeast European-Southwest Asian "superprovince" sensu Bernor et al. (1996) or the Greco-Iranian (i.e. Balkan-Iranian) province sensu Bonis et al. (1992), during that period. ...
Premières découvertes d'autruches (Aves, Struthioniformes, Struthionidae) du Miocène supérieur de Bulgarie et discussion taxonomique et zoogéographique. Deux vestiges du membre postérieur, l'extrémité distale d'un tarsométatarse droit et une phalange proximale du troisième doigt gauche, provenant de deux gisements de la Bulgarie du sud-ouest: Kalimantsi (niveaux du Turolien moyen) et Hadzhidimovo-2 (limite MN 11/12) sont décrits et rapportés à Struthio cf. karatheodoris. Nous présentons une revue des restes fossiles d'autruches du Néogène au Quaternaire, ainsi qu'une discussion sur leurs implications écologique et paléogéographique.
... The climatic conditions then became less arid around 5.3 Ma B.P. (Messinian-Zanclean transition). The occurrence of tralugids and cervids in the latest Miocene deposits in Greece and Macedonia is a strong evidence of increased regional humidity (De Bonis et al., 1992). The latest Miocene deposits in the Eastern Mediterranean recorded also the disappearance of several mammal taxa, such as mastodonts, suids (e.g., Microstonyx), giraffids (e.g., Helladotherium, Samotherium), bovids (e.g., Protoryx, Palaeoryx) and micromammals (e.g., Parapodemus), apparently due to a critical change in ecological conditions (Koufos et al., 2005). ...
The extensional intramontane grabens that formed in southwestern Anatolia in the Late Cenozoic bear the unique tectono-sedimentary, palaeontological and palynological record of a region that underwent rapid transition from the last compressional pulses of the Alpine orogeny to the tectonic phase of orogen collapse and the onset of neotectonic regime. The change in tectonic regime was accompanied by regional climatic changes, recorded by the sedimentary environments of the basins and evidenced by palaeontological and palynological data. The late Early to mid-Middle Miocene was characterized by a warm and humid subtropical climate with densely forested wetlands. The late Middle to Late Miocene witnessed a change to arid climatic conditions, with grass-dominated steppe ecosystems. The Pliocene climate was warm and humid, with savannah-type open habitats.
... The geographic area that presently incorporates the Southern Balkans, Turkey and part of the South peri-Pontic region, formed during late Miocene the western domain of the so called Greco-Iranian biogeographic province (Bonis de et al., 1992). Classical late Miocene European large mammal sites such as Pikermi, Maragheh, Samos, Salonique, Titov-Veles etc. highlight the spatial extent of this area that provided thousands of fossil remains, relatable to several dozen mammal taxa. ...
During the Turolian (late Miocene), the Eastern Mediterranean region is considered to have been part of a single major ecological area, supporting a particular savanna-type large mammal community, referred to as the Pikermian Biome. Analysis of the timing, turnover patterns, biogeographic relations and palaeoecological profile of the Turolian large mammal faunas from either sides of the Aegean Sea failed, however, to confirm the presence of a homogeneous mammal community isotropically behaving through time. For most of the Turolian, the large mammal assemblages from Southern Balkans and Anatolia appear to have existed under different environmental conditions, partly isolated by natural barriers. Overall climatic changes and regional physico-geographic factors, around 7.2 My, allowed the Southern Balkan biogeographic region to be temporarily part of the sub-Paratethyan bioprovince. As a result, significant faunal reorganizations and interchanges triggered the emergence and expansion of the “Pikermian” mammal fauna, which collapsed soon after 7.0 My as a consequence of the early Messinian global changes. The Pikermian Large Mammal Event seems to follow known procedures related with contemporaneous marine and land faunal episodes across the Mediterranean.
... The faunal assemblages from the Late Miocene localities of the Potwar Plateau, in the Siwaliks province, differ greatly from those of the GIA province by a higher diversity of Suina, Cervoidea and Primates. Moreover, the absence in the Potwar Plateau and the presence in Molayan of various taxa (Hyracoidea or Schizotheriinae) comfort the affinities with the Greek, Iranian, and Turkish localities Heintz and Brunet, 1982;Brunet et al., 1984;Bonis et al., 1992). ...
Dental microwear of Late Miocene artiodactyls from Afghanistan can yield information on paleodiet and thus inferences can be drawn about their paleoenvironment. The “Molayan” locality lies at the border between the Greek–Iranian and Siwalik bioprovinces. Knowledge of the paleoenvironment will further our understanding of the faunal exchanges between these two provinces during the Late Miocene. Ninety-nine specimens of seven bovid and one giraffid species are considered. The number and diversity of the samples provide objective data, which enable the reconstruction of the paleoenvironment.Prostrepsiceros aff. vinayaki and Sporadotragus tadzhikistanicus have similar dental microwear pattern, which suggests a “meal by meal” mixed feeding diet. Phronetragus aff. secondus, Gazella sp., and the giraffid Palaeotragus cf. rouenii can only be characterized as mixed feeders in a general sense. Dorcadoxa porrecticornis was an obligate grazer and the two species of Tragoportax were variable grazers. The analysis also brings out clear similarities between extinct grazers and Equus przewalskii, which is a C3 grazer. Isotopic evidence from previous studies supports the microwear data. Comparisons with data from the Late Miocene of the Potwar Plateau suggest strong environmental differences with the Afghan locality. When brought together, this disparate evidence about diet suggests an open and dry environment composed mainly of C3 grasses and evergreen bushes during the Late Miocene in Afghanistan.
In this article, we describe so far unpublished proboscidean specimens from several Late Miocene localities of Romania. A partial mandible and the complete upper/lower cheek tooth rows of a deinothere individual from the site of Gherghești 1 belong to Deinotherium proavum and comprise one of the few examples of entire cheek tooth rows of the same individual of this species. Gherghești 1 is geographically close to Mânzaţi from where the celebrated skeleton of “Deinotherium gigantissimum” was discovered at the end of the nineteenth century, and thus further highlights the importance of Romania in the study of this emblematic deinothere. Deinotherium proavum represents the last deinothere species in Europe and corresponds to the terminal stage of the size increase characterizing the evolution of European deinotheres. Two zygodont molars are attributed to the rare “Mammut” cf. obliquelophus and add to the scarce record of “Mammut” in the Miocene of Eurasia. They document the secure presence of “Mammut” in the Miocene of Romania. The small size of the studied molars compared to known specimens of the Pliocene “Mammut” borsoni and the weak development of the distal cingulum in the lower third molars may have taxonomic and biostratigraphic importance. Furthermore, the presence of an amebelodontid is documented by a large-sized and dorsoventrally flattened lower tusk fragment that shows tubular dentine in its inner part and is attributed to the tetralophodont shovel-tusker Konobelodon. This specimen marks the first record of the genus in Romania. Finally, the biostratigraphic distribution of the taxa is discussed.
Proboscideans (Mammalia: Proboscidea) originated during the Eocene (perhaps already during the Paleocene) in Africa. Their fossil record narrates an amazing evolutionary history, ranging from the Paleogene to the Quaternary. Proboscideans experienced in the past a great diversification and wide distribution in Africa, Europe, Asia, and the Americas. They persist until today with only two genera, Loxodonta and Elephas, geographically confined in regions of Africa and Asia, respectively. The review of the fossil record of the Neogene proboscideans (excluding the members of Elephantidae that are treated elsewhere) in Greece revealed the presence of deinotheres (Deinotheriidae), mammutids (Mammutidae), choerolophodonts (Choerolophodontidae), amebelodonts (Amebelodontidae), tetralophodont gomphotheres (Gomphotheriidae), and stegodonts (Stegodontidae) in more than fifty localities, ranging from the early Miocene to the Early Pleistocene. Fourteen taxa are here considered valid, three of them (Choerolophodon chioticus, C. pentelici, and Konobelodon atticus) erected from type localities in Greece. The most diverse localities are Pikermi and Samos, where at least four proboscidean species have been recorded. The peak in taxonomic diversity occurred during the Turolian (late Miocene). The Greek proboscidean fossil record contains several highlights. The earliest appearance of the family Deinotheriidae in Europe is documented in Gavathas of Lesvos Island, and it is the proboscidean family with the widest temporal distribution in Greece. A deinotheriid skull from Samos Island is so far the most complete juvenile one known from Eurasia and Africa. Choerolophodon presents the widest temporal distribution among the genera in Greece, and where present, it is the dominant genus in terms of abundance. The rich choerolophodont sample allows the distinction into evolutionary stages and renders the genus as biostatigraphically important for Southeastern Europe. The late Miocene Anancus from Chomateri represents the first appearance of the genus in Greece and one of the earliest occurences in Europe. The sample of the late Pliocene Mammut borsoni from Milia, Grevena, is the richest one of this species, including partial skeletons, the longest upper tusks ever recorded in the world and the most complete mandible in Europe. During the Pliocene–Early Pleistocene, the most frequent and widespread proboscidean is the last European gomphothere Anancus arvernensis. Finally, the Siatista Stegodon is the first evidence of the presence of stegodontids in Europe.
Hipparionines represent the greatest part of the Nikiti 2 (NIK) fossil mammal collection. The determination of the material indicates the presence of four distinct species. Two small-sized forms are distinguished in the NIK material: Hipparion macedonicum, well known in northern Greece, and a new small-sized species named Hipparion sithonis. The main character separating the two small-sized species is the presence of a canine fossa in H. sithonis nov. sp., as well as its slightly larger size, deeper narial opening, and the relatively larger and more robust metapodials. The medium-sized NIK hipparion is separated from other known medium-sized forms, representing the new species Hipparion philippus. Finally a large-sized form similar to H. proboscideum is identified in the studied material; as it is only represented by a few specimens, it is referred to as H. cf. proboscideum. The morphological characters of the hipparions, as well as their stratigraphic range suggest an early Turolian to lower middle Turolian (MN 11- lower MN 12) age for the NIK fauna. The relations of the new species, as well as the palaeobiogeography of the Aegean region during the Turolian are discussed on the basis of the hipparions found in NIK and in a wider area.
Two finds, distal end of right tarsometatarsus and a proximal pedal phalanx of the left 3(rd) toe, from two sites (Kalimantsi - the middle Turolian levels, and Hadzhidimovo-2, MN 11/12 boundary, SW Bulgaria) are reported. They are referred to Struthio cf. karatheodoris. The paleoecological and paleozoogeographical aspects of the presence of struthionds are discussed. Struthio remains supported the concept that the openlands were a typical element of the mosaic landscape of the open woodlands/park type forest landscape in the western Pontic part of the Pikermian biome of the Balkan-Iranian late Miocene zoogeographical province.
Both China and Greece have abundant fossils of the late Miocene Hipparion fauna. The habitat of the Hipparion fauna in Greece was a sclerophyllous evergreen woodland. The Chinese late Miocene Hipparion fauna is represented respectively in the Guonigou fauna (MN 9), the Dashengou fauna (MN 10), and the Yangjiashan fauna (MN 11) from Linxia, Gansu, and the Baode fauna (MN 12) from Baode, Shanxi. According to the evidence from lithology, carbon isotopes, paleobotany, taxonomic framework, mammalian diversity and faunal similarity, the paleoenvironment of the Hipparion fauna in China was a subarid open steppe, which is different from that of Greece. The red clay bearing the Hipparion fauna in China is windblown in origin, i.e. eolian deposits. Stable carbon isotopes from tooth enamel and paleosols indicate that C 3, plants dominated the vegetation during the late Miocene in China. Pollens of xerophilous and sub-xerophilous grasses show a signal of steppe or dry grassland. Forest mammals, such as primates and chalicotheres, are absent or scarce, but grassland mammals, such as horses and rhinoceroses, are abundant in the Chinese Hipparion fauna. The species richness of China and Greece exhibits a similar trend with a clear increase from MN 9 to MN 12, but the two regions have low similarities at the species level. The entry of Hipparion led to a comparable radiation in China and Greece, but their dietary evolution is different. In conclusion, the ecosystems of the Chinese and Greek Hipparion faunas have an obvious dissimilarity in the late Miocene. © Publications Scientifiques du Muséum national d'Histoire naturelle.
The determination of the palaeoecological conditions of the Perivolaki fauna is given in the present article using various methods. The faunal diversity of Perivolaki is studied using several indices (Simpson, Shannon-Wiener, Whittaker) and indicates a homogeneous and equilibrated fauna with normal taxa distribution. The dental microwear analysis of the ungulates (Bovidae, Equidae) provided data about the feeding preferences of the identified taxa and in comparison to those from other localities a bushy-shruby-woody palaeoenvironment is possible for Perivolaki. The faunal composition of Perivolaki fauna suggests the dominance of the bovids and equids, indicating a relatively open environment. The comparison of the faunal composition of the Perivolaki fauna with those from various European localities, as well as with the recent ones from certain environments, using multivariate analysis suggests that the Perivolaki fauna matches to the recent open and dry ones. The faunal similarity is also studied using the Simpson's index, indicating close relations of the Perivolaki fauna to those of Axios valley than to those of Southern Greece (Pikermi, Halmyropotamos), Eastern Aegean Sea (Samos) and Asia Minor. All the available results from this study suggest for Perivolaki an open bushy-woody environment with grass undergrowth.
Palaeogeographic and climatic changes in the Eastern Mediterranean during the Neogene/Quatemary led to extended mammalian migrations and faunal exchanges between Eurasia and Africa. At the same time, the Beringian landbridge was activated several times, and American faunal elements entered Eurasia. It appears that the main factor affecting migration potential and faunal changes/exchanges during the Neogene was palaeogeography, while after the early Pliocene migrations were mainly controlled by climatic changes. Several mammalian migrations can be distinguished, but the most important was that of the middle Orleanian at about 17.0-18.0 Ma when Africa and Eurasia were connected after a long separation and a great number of African faunal elements entered Eurasia and vice versa. Some more important faunal changes also occurred: 1. at ∼5.5 Ma, marking the beginning of the Pliocene, 2. at ∼2.0-1.8 Ma, marking the beginning of the Pleistocene, and 3. at ∼1.0 Ma, defining the early/middle Pleistocene boundary and the establishment of modem mammal fauna. During the Pleistocene, oscillation of glacial and inter-glacial periods caused an alternation of cold-steppic faunas with temperate ones in the Eastern Mediterranean. Endemic late Pleistocene mammalian faunas developed in the Mediterranean islands after their isolation ; "dwarf" elephants, cervids and hippos occurred, as well as giant rodents.
We describe here new remains of the giraffid Bramatherium, from the late Miocene-early Pliocene of Hasnot, northern Pakistan. The fossil giraffid material from Hasnot represents 2 taxa of Bramatherium, B. megacephalum and B. grande. The Hasnot locality is characterised by high proportions of ruminants, namely Boselaphini, Antilopini, Sivatheriinae, Cervini, Dorcatherium, and Dorcabune. The Hasnot fossil ruminant assemblage is fairly typical for the Siwalik province and comparisons with the Greco-Iranian-Afghan and African palaeobiological provinces indicate a late Miocene-early Pliocene age. It is suggested that local Hasnot palaeoenvironments comprised wetlands to woodland biomes, perhaps devoid of expansive dense forests.
Different opinions have been expressed on the age of the Molayan mammal locality in Afghanistan which has been estimated as middle or late Turolian. This paper reviews the available biochronological data, mainly provided by affinities of Hipparion molayanense, Mesopithecus pentelicus and rodents, and compares this fauna with those from the Siwaliks of Pakistan and of the Aegean area. These comparisons favor the correlation of Molayan with the middle Turolian (MN12).
Recent systematic excavations near the village of Kerassia (Northern Euboea) have yielded a diverse mammal fauna of Late Miocene age. Among the most complete and prominent findings are a juvenile skull and an adult mandible that belong to two different horned rhinoceros species. The juvenile skull from the site Kerassia-3 (K3) is assigned to Ceratotherium neumayri, while the adult mandible from the plausibly isochronous site Kerassia-4 (K4) belongs to Dihoplus pikermiensis. These new specimens are compared with the known Eastern Mediterranean rhinocerotid record, which is briefly reviewed and updated. The potential interspecific interaction of both species is discussed. Most Eastern Mediterranean localities have yielded only one tandem-horned rhino. In localities with ample material (Pikermi, Samos), where both species are present, one of them is more abundant, signifying a clear interspecific dominance. D. pikermiensis must have preferred temperate forested habitats, whereas the more specialized C. neumayri more open habitats. For the cases of sympatry, a marked resource partitioning is suggested, not excluding some territorial interaction by water resources or at the boundaries of mixed habitats. A partial dietary competition in these eases cannot be excluded, as well.
The fossil Primates (other than Homo)) from the Late Miocene to the Middle Pleistocene found in Italybelong to the superfamilies Hominoidea and Cercopithecoidea. Both have been known since the 19th century. The Hominoids are represented solely by the endemic Oreopithecus bambolii,, the Cercopithecoids by one colobine (Mesopithecus)) and one cercopithecine (Macaca)). A better understanding of these three taxa is expected to be obtained from the study of several recent finds. Oreopithecus bambolii is part of the “Maremmian” fauna peculiar to the Late Miocene in Italy. Most finds have come from the Bacinello lignite mine in the province of Grosseto, where recent discoveries have allowed a direct comparison to be made between specimens from different levels in the sedimentary succession. Mesopithecus is a small to medium-sized Late Miocene to Middle Pliocene colobine represented by two species in both Southern and Central Europe: M. pentelicus occurs in four Italian Miocene sites, whereas the younger M. monspessulanus for the moment is confined to Villafranca d'Asti (Early Villafranchian), where a new specimen has recently been discovered. Previously Macaca fossils from the Middle Pliocene to the Middle Pleistocene of Italy have been divided into two species: M. florentina,, which is probably related to the living M. sylvanus,, and M. majori,, an endemic dwarf species from Sardinia. The latest finds are those of 8 individuals, provisionally regarded as belonging to the M. florentina-sylvanus lineage, and discovered in association with a Late Villafranchian (Early Pleistocene) vertebrate fauna in a lignite mine at Pietrafitta, Province of Perugia. This welldated collection comprises both gnathic and postcranial fragments and should serve to provide a major contribution towards the understanding of the still problematical taxonomy, phylogeny and palaeoecology of this species.
During his nearly 50-year long career Louis de Bonis has positively and markedly influenced many generations of paleontologists. In this introduction to the volume celebrating the 75th birthday of this French outstanding scientist, we present and discuss the significance of his many contributions to the study of mammalian evolution, focusing especially on his works on primate and carnivoran systematics and evolution, and on his involvement in field excavations in northern Greece Ouranopithecus-bearing sites.
The biogeographic and biodiversity evolution of the European Neogene ungulate faunas is herein addressed by means of faunal similarity and diversity estimators. From the early to the early late Miocene, faunal ressemblance, familial diversity and faunal homogeneity between the various regions increased. They were followed by a progressive decline of diversity towards Pliocene times. We discuss the biodiversity and faunal composition evolution within four bioprovinces and show that in a global context of climate relative stability in the early and middle Miocene, particular palaeogeographic configurations allowed the immigration of a large number of African and Asian families, increasing biodiversity. The late middle to early late Miocene period is characterised by a global climate change (cooling and aridification) that profoundly affects the European ungulate faunas. Indeed, “modern” ruminants (bovids, cervids, giraffids) become the dominant groups of ungulates by the early Pliocene. This dominance is associated with a decrease in familial diversity and with a modern pattern of latitudinal differentiation, with mainly bovids colonising southern regions and, in a lesser extent, cervids developping in northern areas.
Although Molayan is the richest fossil mammal locality in Afghanistan, only a few species have been studied. This paper describes three rodents (Hystrix aryanensis sp. nov., Parapeolomys sp. and Pseudomeriones latidens sp. nov.) and one insectivore (Erinaceus sp.). Lithostratigraphical correlations and biostratigraphical studies in the Khurdkabul Basin provide evidence that the Molayan locality is younger than three other localities in this basin: Sherullah, Taghar and Ghazgay. In addition, the taxa described here, along with several species of large mammals from Molayan, permit correlations with the Siwalik succession in Pakistan to the east, and with European Neogene mammal ages to the west. The age of the Molayan fauna is concluded to be mid Turolian. Observations on the mammal faunas from the Khurdkabul Basin suggest that the altitude of the area, at present over 2000 m, was much lower during the late Miocene. Uplift of the basin probably occurred no earlier than the ?late Pliocene.
This paper describes the first fossil porcupine remains from Iran. Four upper cheek teeth and two fragmentary lower incisors
present sufficient characters for identification as Hystrix aryanensis, a species previously known from the late Miocene locality of Molayan (Afghanistan) estimated at ca. 7–8Ma. The dental features
of porcupines are discussed to show their systematic value and highlight evolutionary trends in late Miocene and Pliocene
porcupines. This study also discusses the dispersal history of fossil porcupines in relation to paleobiogeographic provinces
and environmental changes during late Miocene to late Pliocene time.
Diese Arbeit beschreibt die ersten Stachelschweinfossilfunde aus dem Iran. Vier Oberkieferzähne und zwei fragmentäre untere
Frontzähne zeigen Merkmale, die zur Bestimmung von Hystrix aryanensis ausreichen, eine Art, die bisher aus dem jüngeren Miozän aus Molayan (Afghanistan) bekannt ist und auf ca. 7 bis 8Ma geschätzt
wird. Die Zahnmerkmale der Stachelschweine werden diskutiert, um deren systematische Aussagekraft hervorzuheben, und um die
Evolutionstrends der Stachelschweine im jüngeren Miozän und Pliozän zu beleuchten. Diese Arbeit diskutiert auch die Entwicklung
der Verteilung von fossilen Stachelschweinen in Verbindung mit den paläobiogeographischen Provinzen und Umweltveränderungen
während des jüngeren Miozäns und Pliozäns.
KeywordsHystricidae-Rodentia-Late Miocene-Iran-Biogeography
SchlüsselwörterHystricidae-Rodentia-Jüngeres Miozän-Iran-Biogeographie
The new mammal locality of Akkasdagi yielded a rich fauna dated as Middle Turolian. It is situated in the southern part of the Çankiri-Çorum Basin in central Anatolia. In the Akkasdagi area, the sedimentary deposits, mapped as the Kizilirmak Formation, consist of tuffs and lacustrine limestones intercalated with clastic deposits, whilst in its type section situated to the north near Kizilirmak town, this formation is mainly formed of fluviatile deposits. Facies distribution and geological observations led to conclude that during the Miocene, depositional centers migrated from the north to the south of this basin.RésuméLe nouveau gisement de mammifères à Akkasdagi, au sud du bassin de Çankiri-Çorum (Anatolie centrale), a livré une abondante faune datée du Turolien moyen. Dans le secteur d'Akkasdagi, la série sédimentaire attribuée à la formation de Kizilirmak comporte une succession de tufs et de calcaires lacustres, intercalés dans les sédiments détritiques. Cette succession diffère de celle de la coupe type, située plus au nord, qui est dominée par des dépôts fluviatiles. Les observations géologiques montrent qu'au cours du Miocène, le centre de sédimentation a migré du nord vers le sud.
More than 120 rhinocerotid remains unearthed in the middle Turolian locality of Akkaşdaǧι (Central Anatolia) are described. The fauna is diversified, with a pair of large two-horned species, Ceratotherium neumayri (Osborn, 1900) and Stephanorhinus pikermiensis (Toula, 1906), and two smaller species of short limbed aceratheriines (Chilotherium sp. and Acerorhinus sp.). Ceratotherium neumayri is by far the most common species, with a complete skull, 114 specimens and at least 11 individuals. The cranial, dental and postcranial remains of C. neumayri are among the largest ones described so far for this species. The coexistence of C. neumayri, S. pikermiensis, and chilotheres is common in the Turolian of Eastern Mediterranean: comparable rhinocerotid associations are known at Kavakdere (MN 12, Turkey) and Samos (MN 12, Greece). The large size of the C. neumayri specimens is consistent with the middle Turolian age for Akkaşdaǧι (MN 12), as stated on the whole mammalian fauna and radiometric data. © Publications Scientifiques du Muséum national d'Histoire naturelle, Paris.
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