Pliocene vertebrate locality of Qalta, Ankara, Turkey. 12. Paleoecology, biogeography and biochronology

Sevket SEN Laboratoire de Paléontologie, URA 12 du CNRS, Muséum national d'Histoire naturelle, 8 rue de Buffon, F-75231 Paris cedex 05 (France) [email protected]

Geneviève BOUVRAIN Laboratoire de Paléontologie des Vertébrés, URA 1761 du CNRS, université Pierre-et-Marie-Curie, 4 place Jussieu, F-75252 Paris cedex 05 (France)

Denis GERAADS EP 1781 du CNRS, Collège de France, 30 avenue Marcelin-Berthelot, F-92360 Meudon (France)

Sen S., Bouvrain G. & Geraads D. 1998. — Paleoecology, biogeography and biochrono­ logy, in Sen S. (ed.), Pliocene vertebrate locality of Qalta, Ankara, Turkey. 12, Geodiversitas 20 (3) : 497-510.

ABSTRACT Integrated studies on the Pliocene vertebrate locality of Calta provided abun­ dant data on the environment of the fauna, its biogeographic relationships and its age. The fauna is dominated by animals adapted to relatively dry and KEYWORDS open environments with loose soil and some watet points. The presence of Mammals, short-limbed herbivores and of ochotonids might indicate a landscape with Pliocene, a ta Ruscinian, some relief nearby. The C l fauna is mainly composed of elements with Turkey, European and some Asiatic affinities. African affinities of some taxa are paleoenvironment, doubtful. The time range of some mammalian groups is discussed, the Caha biogeography, biochronology. locality is dated as eatly MN15 (late Ruscinian).

1998 • 20 (3) 497 Sen S., Bouvrain G. & Geraads D.

RÉSUMÉ Le gisement de vertébrés pliocenes de Çalta, Ankara, Turquie. 12. biogéographie et biochronologie. Des recherches multidisciplinaires sur la faune de vertébrés et les sédiments de Çalta ont permis d'obtenir des résultats sur l'environnement, les affinités de la faune et son âge. Cette faune est dominée par des éléments vivant dans un milieu telativement sec et ouvert, avec cependant quelques points d'eau et un sol meuble. Quelques hetbivotes à

MOTS CLES pattes couttes et l'abondance des Ochotonidae suggèrent des reliefs dans le Mammifères, paysage. Dans cette faune, les groupes à affinités européennes sont plus nom­ Pliocène, Ruscinien, breux que ceux à affinités asiatiques. Les rapports africains de quelques Turquie, taxons sont sujet à caution. La synthèse des données biochronologiques per­ paléoenvironnemenr, met de corréler ce gisement à la partie inférieure de la zone MN15 biogéographie, biochronologie. (Ruscinien supérieur).

INTRODUCTION Reptiles Testudo sp. In the present monograph on the Caha vertebrate Scincidae indet. fauna, the sedimentology of deposits and the iso- Lacertidae indet. topic values of bones, teeth and soil samples are Ophisaurus sp. studied. Previous papers reported on amphibians, Varanus marathonensis Weithofer, 1888 reptiles and some mammals, and also on the geo­ Amphisbaenidae indet. logical context of this locality. Thetefore, a com­ Scolecophidia indet. plete and reliable faunal list of Ca'ta is now Eryx sp. available. Colubtidae indet. All these studies provide numerous data on envi­ Palaeonaja sp. ronmental habits of various taxa and their bio- Aves geographic affinities. Moreover, the new criteria Struttolo sp. allow a better estimate of the age of this locality. Consequently, it is now possible to provide a Mammalia synthetic view on the paleoecology of the site, Insectivora the resemblances of the fauna with its equivalents Asoriculus gibberodon (Petenyi, 1864) from other regions, and to discuss its age. Mafia csarnotense Reumer, 1984 Soricidae indet. Erinaceus sp. FAUNAL LIST Rodentia *Centralomys magnus (Sen, 1977) a The faunal list of the C ha locality is as follows Occitanomys sp. a (the new taxa described from C ha are indicated *Orientalomys galaticus (Sen, 1975) by an asterisk): Apodemus dominans Kretzoi, 1959 Mesocricetus sp. cf. M. primitivus de Bruijn et Gastropoda al, 1970 Helix cineta Müller, 1781 *Pseudomeriones tchaltaensis Sen, 1977 Subzebrinus sp. Mimomys davakosi van de Weerd, 1979 Amphibians Pliospalax macoveii (Simionescu, 1930) Pelobates sp. Dryomimus eliomyoides Kretzoi, 1959 Bufo sp. cf. B. viridis Laurenti, 1768 Lagomorpha Rana sp. *Ochotonoma anatolica Sen, 1998

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Carnivora Rhinocerotidae *Vulpes galaticus Ginsburg, 1998 Nyctereutes donnezani (Depéret, 1890) *Chasmaporthetes kani anatolicus Ginsburg, 1998 Lynx issiodorensis (Croizet et Jobert, 1828) Machairodus giganteus (Wagner, 1848) Perissodactyla Dicerorhinus megarhinus (de Christol, 1834) *Hipparion heintzi Eisenmann et Sondaar, 1998 Hipparion sp. cf. H. longipes Gromova, 1952 Artiodactyla Gerbillidae A Sus arvernensis minor Depéret, 1 890 Cervus sp. cf. C. perrieri Croizet et Jobert, 1828 Giraffa sp. cf. G. jumae Leakey, 1965 *Gazella emilii Bouvrain, 1998 *Tchaltacerus longicornis Bouvrain, 1998 Bovidae indet. cf. Koufotragus bailloudi (Arambourg et Piveteau, 1929) Gazellospira sp. Hippotragini indet.

PALEOENVIRONMENTAL RECONSTRUCTION

The community of vertebrates from Calta reflects quite well the environment in which it lived; this environment seems to have been contrasted, juxtaposing different types of bio- topes. The pie-diagram of figure 1A gives the relative proportions (in terms of MNI) of the various groups present at Calta. Systematic screening yielded a great number of micromammals, among which gerbils are dominant, thus poin­ ting to an open environment. When only large mammals are considered (Fig. IB), we may notice the abundance of carnivores, probably greater Fig. 1. — Mammalian faunal composition of Calta and than in living assemblages. As in the late Kemiklitepe A + B localities based on the minimum number of individuals. A, Calta mammalian fauna with all groups; B, the Miocene of Kemiklitepe A + B (Fig. 1C, Bonis et same fauna without small mammals; C, Kemiklitepe A + B fauna al. 1994), bovids are the dominant group among without rodents. artiodactyls, but the latter are more diverse at Calta, with the addition of a suid and a cervid, missing at Calta; some of them (Tubulidentata, than at Kemiklitepe A + B. However, several Hyracoidea, Chalicotheriidae) may have become groups which occur in the upper Miocene are extinct in the meantime, but proboscideans sur-

GEODIVERSITAS • 1998 • 20(3) 499 Sen S-, Bouvrain G. & Geraads D.

vived in the area until the Pleistocene, and their burrows (Sen this volume). The pig Sus arver- absence here might have an ecological meaning. nensis minor is also known as a inhabitant of We may also, once more, notice the lack of cer- humid and hot loose soils in which it looks for copithecid primates, a group which remains its food (Guérin et al. this volume). completely unknown in Turkey. Two groups clearly indicate the presence of water bodies: frogs, although they are not very abun­ dant, and an aquatic shrew, Asoriculus, which FUNCTIONAL ADAPTATIONS represents 64.9% of the insectivore remains One of the characteristics of this fauna is the (Reumer this volume). The presence of some marked lengthening of the distal segment of limb murids (Sen this volume) and a deer (Geraads bones in several herbivores. This lengthening is this volume) evidences the existence of a zone an adaptation to running and implies therefore with abundant vegetation cover in the area. the presence of open landscapes. Thus Hipparion The landscape of Çalta, when the fossiliferous cf. longipes, as its name indicates, is characterized deposits accumulated, seems to be mainly com­ by lengthened limbs (Eisenmann & Sondaar this posed of steppic environments (cursorial ani­ volume); Gazella emilii has limb bones longet mals, strong percentage of gerbils, spalacids and than those of extant gazelles, and this is also the ochotonids, Sen this volume), with some water case of the Hippotragini (Bouvrain this volume). sources (persistent or temporary?), forested zones Dicerorhinus megarhinus is also a cursorial rhino and also probably with some relief. Such an envi­ (Guerin & Sen this volume). The Calta giraffe is ronment is in agteement with sedimentological close to the African Plio-Pleistocene Giraffa results (Lunkka et al. this volume). jumae, which allows us to envisage a similar bio- The climate could have been dry. Indeed the tope to that of the living giraffe (Geraads this weak specific diversity of Soricidae would imply volume) .This is also the case for the Calta ostrich a dry and even arid climate (Reumer this volu­ (Janoo & Sen this volume). me). The specialization of the nasal region of Hipparion heintzi recalls the disposition existing However, beside these herbivores with lengthe­ in the Saïga antelope or in Camelidae and would ned limb bones, there exist two taxa showing a mean an adaptation to an arid climate different adaptation: Tchaltacerus is a small ante­ (Eisenmann & Sondaar this volume). Isotopic lope with short and robust metapodials; such analyses show on the one hand the absence of C4 metapodials are currently known in bovids living rich plants, therefore similar to the ptesent day in steep landscapes such as caprines (Bouvrain situation in central Anatolia, and on the other this volume). The strong shortening of the meta­ hand a strong évapotranspiration of plants cor­ carpal bones in Hipparion heintzi as compared to responding to an open environment with arid the radius is found only in mountain zebras; conditions (Bocherens & Sen this volume). however the widening of the hoofs in Hipparion Independently of indications provided by each heintzi contradict an adaptation to a steep land­ taxon, one can try to draw conclusions from the scape and would rather imply the presence of global analysis of the Çalta mammalian commu­ soft ground (Eisenmann & Sondaar this volu­ nity. From the various possible approaches, we me). The presence of an ochotonid also indicates have retained the method of the cenograms and the presence of mountain meadows and rocky the factor analysis of the taxonomic composition. grounds (Sen this volume). Another characteristic of the Calta vertebrate fauna is the great number of animals having a CENOGRAM burrowing mode of life or even living under­ The cenogram (Valverde 1964; Legendre 1986) ground. It is the case of several amphibians and is a graph established by plotting the species on reptiles: Pelobates, Eryx, amphisbaenians and sco- the X-axis in order of decreasing weight against lecophidians (Rage & Sen 1976), but also of the natural logarithm of their weight (in grams) some rodents such as the Spalacidae and even the on the Y-axis. The slope of the line(s) joining the Gerbillidae that dig shallow but labyrinthine points that correspond to the weight of each

500 GEODIVERSITAS • 1998 • 20 (3) Pliocene paleoecology, biogeography and biochronology of Calta

16.,

• 14- I • * * CALTA 12 J • •

•OJ • • ^

* • • • .

10 15 20 25 30

LOKORI LA CALERA LAYNA PERPIGNAN

FIG. 2. — Cenogram of the Qalta mammalian fauna, compared with those of Lokori (present day dry woodland of Kenya), and of three Pliocene sites of western Europe. The data for the western European Pliocene sites is taken from Montuire (1994). The X and Y axes scales given for the Qalta cenogram are also used for the others. taxon and the presence of possible breaks along open forest/wooded grassland environment with this line(s) provide, according to these authors, a subarid climate. On the Calta cenogtam, the indications on the humidity of the environment, slope of larger mammals is not much steeper its degtee of openness, and even a minimal tem­ than that of smaller ones, implying that the cli­ perature estimation. We believe, however, that mate was not very arid, less so than, e.g., that of these graphs must be interpreted with caution, La Calera (MN14, Spain). The cenograms of the because the slope of the curve depends first on Spanish and French sites mentioned above are the number of species, therefore on the impor­ taken from Montuire (1994). tance of excavations, and second on the graphic scale used. The most noticeable feature of the TAXONOMIC COMPOSITION Calta cenogram (Fig. 2) is the wide gap between Mammalian communities also differ by their larger (more than 18 kg) and smaller (less than taxonomic composition, since the main mamma­ 0.5 kg) species. This would suggest an open lian groups are more or less abundantly represen­ environment, much more open than those of the ted depending on environmental factors. toughly contemporaneous sites of Layna Multivariate analyses allow a better estimation of (MN15, Spain) and Perpignan (MN15, France), these differences. We have thus undertaken a fac­ and probably slightly more open than that of the tor analysis on twenty extant communities of the extant Lokori fauna in Kenya which is from an Ancient World (Table 1), by using as variables

GEODIVERSITAS • 1998 • 20(3) 501 Sen S., Bouvrain G. & Geraads D.

-1J

X Taxa • Recent faunas; forest • Fossil faunas; Africa

O Recent faunas; savanna • Fossil faunas; Eurasia

FIG. 3. — Correspondance factor analysis of Recent and Pliocene mammalian faunas. Matrix from references cited in tables 1, 2.

TABLE 1. — List of the sites with living mammalian communities used for factor analysis. Their environments and the related refe­ rences are also given.

Sites Countries Environment Références

Umfolozi A South Africa savanna with acacias, gallery forest Mentis 1970 Hluhluwe B South Africa decidual humid forest Bourquin et al. 1971 Chobe C Botswana flood plain Sheppe & Haas 1976 Kafue D Zambia flood plain Sheppe & Osborne 1971 Baoulé E Mali dry savanna Bousquet 1992 Lomto savanna F Ivory Coast savanna with acacias, gallery forest Bourlière et al. 1974 Niokolo-Koba G Senegal wooded savanna, flood plain Bousquet 1992 La Comoé H Ivory Coast wooded savanna, flood plain, forest Lartigues & Poilecot 1997 Azagny J Ivory Coast marsh and forest Bousquet 1992 Tai K Ivory Coast evergreen tropical forest Bousquet 1992 Lamto forest L Ivory Coast montane forest Bourlière et al. 1974 Makokou M Gabon equatorial forest Happold 1996 Irangi N Congo equatorial forest Rahm 1996 Mount Kivu O Former Zaire montane forest, savanna, marsh Rahm & Christiaensen 1963 Gunong Benom P Malaysia tropical rain forest Medway 1972 Kuala Lumpat Q Malaysia tropical rain forest Medway & Wells 1971 Gunong Mulu R Malaysia tropical rain forest Anderson et al. 1982 Wilpattu S Sri Lanka forest, flood plain Eisenberg & Lockhart 1972 Gir T India open forest Bernick 1974 Kan h a u India forest, meadow Schaller 1967

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TABLE 2. — List, age and references of Pliocene localities used for factor analysis.

Localities Countries Age References

Layna 1 Spain MN15 Perez & Soria 1989-1990 La Calera 2 Spain MN14 Alcalá 1994 Villaroya 3 Spain MN16 Azanza et al. 1989 Serrat-d'en-Vacquer 4 France MN15 Mein 1990 Les Etouaires 5 France MN16 Heintz et al. 1974 Dorkovo 6 Bulgaria MN14 Thomas et al. 1986 Malusteni 7 Romania MN15 Macarovici 1974 Kvabebi 8 Georgia MN16 Vekua1972 Karboliya 9 Moldavia MN14/15 Pevzner et al. 1996 Yushe (Upper Gaozhuang) 10 China MN15 Tedford et al. 1991, Fejfar et al. 1997 Calta CALTA Turkey MN15 this volume Dinar-Akgakdy 11 Turkey MN14 Van der Meulen & Kolfschoten 1986 Giilyazi 12 Turkey MN16 Sickenberg et al. 1975 Warwire 13 Uganda 3.5 Ma Pickford & Senut 1994 Nkondo 14 Uganda 4.5 Ma Pickford & Senut 1994 KT 13 15 Tchad 3-3.5 Ma Brunet et a/.1997 Laetoli 16 Tanzania 3.5 Ma Harris et al. 1989 Tulu Bor 17 Kenya 3.5 Ma Feibel et al. 1991 Lokochot 18 Kenya 3.5 Ma Feibel et al. 1991

the main groups: Primata (PRIM), Lagomotpha possible to deduce some paleoecological conclu­ (LAGO), Tubulidentata (TUBUL), Proboscidea sions from their comparison. African fossils fau­ (PROBO), Hyracoidea ((HYRA), Tapiridae nas would thus illustrate less varied habitats than (TAPIR), Rhinocerotidae (RHINO), Equidae Eurasiatic faunas, since they are more grouped (EQUID), Suidae (SUID), Hippopotamidae and indicate in general more open environments. (HIPPO), Cervidae (CERV), Giraffidae This corresponds well with what is known about (GIRAF) and Bovidae (BOVID). We have there­ them. One can then interpret the position of fore excluded carnivores (that yield little ecologi­ Çalta, on the very left of the graph, as reflecting cal information, and whose number depends on a more open habitat than any other Eurasian the type of deposits), as well as bats and rodents locality. This result, even if it does not have to be (except Hystricidae: HYST) whose abundance accepted without nuances, corresponds well to also depends on taphonomic conditions. This indications provided by othets approaches. analysis distinguishes well (plane 1-2, the CFA, Fig. 3) open habitats from closed ones (axis 1 especially), and also African faunas from those of BIOGEOGRAPHY Eurasia. If we replace on the same graph (as sup­ plementary or illustrative elements) a set of fau­ The Çalta fauna is composed of three species of nas from the Pliocene of the Old World amphibians, ten teptiles, one bird and thirty-one (Table 2), one can make several observations. species of mammals. This number clearly shows As previously noticed by de Bonis et ill. (1994), that this association is tich enough to be repre­ fossil faunas are systematically shifted in relation sentative of the tetrestrial faunal spectrum of the to the living ones. The distinction between area during the time of deposition of the fossili- African and Eurasiatic faunas nevettheless ferous hotizon. remains clear, since they are clearly separated in Taken one by one, the genera and species reco­ two different parts of this diagram. One can the­ gnized at Çalta have different affinities when refore rightfully think that the observations on compared to the geographic distribution of rela­ the spatial distribution of extant faunas should ted groups. We will classify them in four catego­ be applied to fossil ones, and that it is therefore ries, having in mind the affinity of each taxon, as

GEODIVERSITAS • 1998 • 20(3) 503 Sen S., Bouvrain G. & Geraads D.

European, Eurasiatic, African and local. Fejfar etal. (1990) and Bachelet (1990) maintai­ However, as we will see below, some taxa found ned it as a distinct species. Later representatives at Calta cannot be clearly related to a biogeogra- of the genus Mimomys are the common elements phic province. For instance, Giraffa may appeal of the late Pliocene and early Pleistocene faunas as an African element of this fauna because of the all over Eurasia. past and present dispersal of all representatives of The Calta rhino Dicerorhinus megarhinus is also the genus, from the latest Miocene to Present; exclusively European. It was discovered in almost however, it may be considered as an Eurasian all countries from Spain to Ukraine and Georgia form if we take into account its phylogenetic during the Ruscinian (Guerin & Sen this volu­ relationships with the genus Bohlinia (late me). The geographic distribution of Sus arver- Miocene, Greece, Iraq). nensis closely follows that of this rhino (Guerin et al. this volume). EUROPEAN AFFINITIES The large overall size, the robustness, and even Among the Calta mammals, the taxa having the upper cheek tooth morphology are rather European affinities are dominant. The insecti- similar in Hipparion heintzi from Calta and in vores Asoriculus gibberodon and Mafia csarnotense H. crassum (Eisenmann & Sondaar this volume). are the common elements of central European The latter species, the type locality of which is Pliocene faunas (Reumer this volume). A. gibbe­ Perpignan in southern France, was also found in rodon was also found in the localities of Greece. Maramena, Maritsa and Tourkobounia-1 and, An indeterminate bovid from Calta can only be M. csarnotense at Apolakkia (Greece). These two compared with Koufotragus bailloudi from the species are not known in western and eastern late Ruscinian of northern Greece. Europe. The European rodents of Calta are Centralomys, Occitanomys, Dryomimus and ASIATIC OR EURASIATIC AFFINITIES Mimomys davakosi. The species included in Among the Calta mammals several groups have a Centralomys were described from Italy, Greece wide geographic distribution across Eurasia. This and Turkey, and this genus has phylogenetic rela­ is the case of Erinaceus sp. and Apodemus domi- tionships with the western European Castillomys, nans which are known in many localities in and perhaps with Occitanomys (see Martin Suarez Eurasia. Orientalomys is a murid which probably & Mein 1991). Occitanomys is a well-known originated from Asia, but also occurred in south­ western European genus, but de Bruijn (1976) eastern Europe and Anatolia. Pseudomeriones has and Black et al. (1980) referred to it, with a ques­ a similar distribution; it was found in China, tion matk, a new species from Turolian localities Turkmenistan, Afghanistan, Turkey and Greece in Greece. Moreover, Storch (1987) included in (see Sen this volume); it was also recorded in the this genus the doubtful "Stepbanomys ?" pusillus Czeck Republic with one tooth, and with one Schaub, 1938 from some latest Miocene locali­ other tooth in Spain! Because of the lack of data ties in Inner Mongolia. If this attribution is cor- on the phylogenetic relationships of this genus, it rect, Occitanomys might have an Eurasian is not possible to recognize from where it origi­ distribution, at least during the late Miocene. nated. Dryomimus is apparently derived from the wes­ The only lagomorph from Calta is a new taxon, tern European Peridyromys or Myomimus (Daams Ochotonoma anatolica, which is related to ochoto- 1981). It occurs during the Pliocene in central nid genera Proochotona, Pliolagomys, Ochoto- and southeastern Europe. Mimomys davakosi is noides and Ochotona. This group has a long one of the oldest species referred to this genus, history and a wide distribution in Asia and eas­ and it was found in localities close in age to the tern Europe; exceptionally its representatives emi­ MN14/MN15 boundary in Greece, France and grated to centtal and western Europe during the Spain. Although Radulescu & Samson (1989) Pliocene and Pleistocene. This group is probably suggested its synonymy with Mimomys moldavi- derived from the early-middle Miocene Bellatona cus Kormos, 1932 from Malusteni in Romania, which is well-known in central and eastern Asia.

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Vulpes galaticus from Calta is the oldest represen­ eastern European localities during the Miocene tative of the genus. Late Pliocene and early and Pliocene, while the extant distribution of Pleistocene occurrences of Vulpes were mentio­ this genus is limited to Africa. The same is also ned all over Eurasia and North Africa. However, true for Giraffa sp. cf. G. jumae. Giraffids are the attribution of several findings from Europe well-known during the middle and late Miocene and Asia to the genus Vulpes is doubtful, for ins­ all over Eurasia and North Africa, but become tance Vulpes odessanus from the Odessa- tare during the Pliocene. The genus Giraffa first Catacombs, and inversely some other records appears in East and South Africa around the described as Cants or Nyctereutes might well refer Mio-Pliocene boundary, and it is quite well- to Vulpes. The other carnivore Lynx issiodorensis represented in the Plio-Pleistocene East African has a geographic distribution covering all faunas. However, this genus is probably derived Mediterranean Europe, the Black Sea coastal from Bohlinia which is only known in late areas and Notth Africa. The genera Nyctereutes, Miocene localities in Greece and Iraq. The Caha Chasmaporthetes and Machairodus have a wide giraffe cannot be the ancestor of its African rela­ geographic distribution from Spain to China and tives since this genus occurs earlier in Africa. to Africa. Although all Calta carnivores have However, it cannot be fully considered as of their relatives in some North African Plio- African origin because of the phylogenetic link Pleistocene localities, they are considered as between this genus and Bohlinia. Thus the Caha having originated from Eurasia or even North gitaffe might be either of eastern European or America (Canidae) and they immigrated to African origin, and we think it would be unwise Africa duting the Pliocene. to draw any biogeographic conclusion from it. The remains of Hippotragini are quite rare at Hipparion cf. longipes is represented at Calta by Calta. It is apparently the unique element of this ten postcranial bones, and is thus rarer than the fauna which originated from Africa, since the other species H. heintzi (Eisenmann & Sondaar Hippotragini first appears in East Africa during this volume). H. longipes is only known from the the late Miocene and later in Europe (Kvabebi, latest Miocene localities of Pavlodar and Georgia) and in the Siwaliks of Pakistan during Kalmakpai in Kazakhstan. However, some rela­ the late Pliocene, about 2-3 Ma ago. ted forms, similar in the proportions of limb Pickford & Morales (1994) demonstrated that bones but slightly smaller, have been found in the entichment of African elements in Spanish the localities of Karaburun in Greece, Layna and Neogene faunas is closely related to climatic fluc­ La Gloria 4 in Spain. tuations, itself regulated by the northern exten- a Gazella emilii from C ha is clearly different from tion of the Ethiopian biogeographic realm. the European Pliocene species G. borbonica, but According to their results, the peaks of faunal quite similar to some species reported from cen­ change, which correspond to the maximum tral Asia (Bouvrain this volume). The new genus extension of the Ethiopian biogeographic realm, and species Tchaltacerus longicornis is a Caprinae occurred at about 7 ± 0.5 Ma and 2.5 ± 0.5 Ma. having resemblances with some species included The rarity of African elements in the Caha fauna in Protoryx from Spain and Georgia. It is in any may be explained by the fact that it belongs to an case related to the Eurasiatic goat-like bovids. intermediate phase between these dates. a ta Some scarce remains of a C ' bovid were deter­ Nevertheless, the Pliocene faunas from the eas­ mined as ? Gazellospira sp. This genus has a wide tern Mediterranean regions are still poorly distribution from Spain to China. known to document enough the phase of African influences in mammalian communities. APRICAN AFFINITIES Taxa with African affinities are rare at Calta- Moreover, if even their living relatives are exclusi­ LOCAL FORMS vely African, they did not necessarily originate Two rodent genera from Calta, Mesocricetus and from this continent. This is the case of the ostrich Pliospalax, are local forms. Their past and present Struthio which is recorded in many Asian and distribution does not exceed the regions surroun-

GEODIVERSITAS • 1998 • 20(3) 505 Sen S., Bouvrain G. & Geraads D.

ding Turkey, i.e. from the Balkans to Iran and some of them are more restricted in time and can the Middle East. be used as key elements to date this fauna. This In summary, the Calta fauna is mainly composed is the case of the insectivore Mafia csarnotense; its of elements with European affinities, togethet oldest known occurrence is at Podlesice (late with some forms that are clearly related to Asiatic MN14, Poland), and the youngest one at groups. Species with African affinities are rare Csarnota-2 (late MN15, Hungary). This species and theit biogeographic relations are doubtful. was also found in Apolakkia (late MN15, Greece). Moteover, studying the Calta insecti- vores, Reumer (this volume) noted the absence BIOCHRONOLOGY of Crocidurinae in this fauna as a proof of its ear­ lier age in comparison to localities whete this The Calta mammal locality is in fluviatile depo­ group exists. Ctocidures originated from Africa; sits which do not contain any volcanic or volca- theit earliest occurrence in Europe is in the loca­ niclastic intercalations. Thus, it is not possible to lity of Apolakkia, Rhodes Island, Greece (Weerd apply radiometric dating technics. In 1994 and et al. 1982; Reumer 1984). This locality is dated 1995, J. Kappelman, J. P. Lunkka, D. Ekart and as late MN15. Later on this group becomes com­ S. Sen sampled a 195 m thick section across the mon in Villanyian and younger faunas in fossiliferous horizon for a magnetostratigtaphic Europe. study. Paleomagnetic analyses of samples have Pseudomeriones is a gerbil which occurs from the been performed by J. Kappelman in the latest Vallesian to ? middle Villafranchian in University of Texas at Austin. Unfortunately, it China, Turkmenistan, Afghanistan, Greece and appeared that most samples are remagnetized Turkey. In the Aegean area, its representatives are probably because of tectonic tilting of deposits P. cf. abbreviatus in latest Vallesian and Turolian after deposition. localities, P. rhodius at Ano Metochi (MN13) The Calta section is not stratigraphically correla­ and Maritsa (MN14) and P. tchaltaensis at Calta. ted to other mammal localities of the basin. The latter species is more derived than the others Moreover, the nearest mammal localities (Sen this volume), which favours an age younger Kavakdere and Coban Pinar yielded Turolian than Maritsa. mammal faunas, and thus they do not contain Mimomys davakosi was only reported from the even one taxon in common with Calta. early MN15 faunas: Ptolemais 3 (type locality, In Turkey, several Pliocene mammal localities are Greece) and Serrat-d'en-Vacquer (reference loca­ already known. However, most of them have pre­ lity of MN15, southern France). It is considered liminary faunal lists, and the others are low in as one of the oldest and most primitive species of species diversity. So, unfortunately, they are not the genus. The great agreement among the reliable for detailed biochronologic comparisons. rodent specialists is that the appearance of In Greece, Pliocene mammal faunas are better Mimomys defines the base of MN15. The other known from a dozen localities (Van der Meulen primitive species of Mimomys is M. antiquus & Kolfschoten 1986; Koufos & Kostopoulos from Peshniovo-Borki and Borehole No. 2 along 1997). Thanks to faunal lists published by these the River Ob in western Siberia (Zazhigin 1980), authors and the studies mentioned therein, the both localities dated as MN15a (Fejfar et al. Calta fauna can be compared with some of them. 1997). Radulesco & Samson (1989) suggested In addition, the close biogeographic affinities of the synonymy of M. davakosi with another pri­ this fauna with those from Eurasia also provides mitive Mimomys from Malusteni (MN15a, biochronologic criteria for correlation. Romania), M. moldavicus Kotmos, 1930. Fejfat Figure 4 shows the biochronologic distribution et al. (1990, 1997) retained this opinion with of some selected genera and species from Calta. some caution because of the poor state of preser­ Most of them appear during the late Turolian or vation of the Malusteni specimens. M. vander- early Ruscinian, and they have a time range meuleni Fejfar et al., 1990 from Spanish localities generally covering the entire Pliocene. However, is also characterized by low dental tracts, strong

506 GEODIVERSITAS • 1998 • 20(3) Pliocene paleoecology, biogeography and biochronology of Calta

VALLESIAN TUROLIAN RUSCINIAN VILLANYIAN E. PLEISTOCENE ^\MN zones

MN9 MN10 MN11 MN12 MN13 MN14 MN15 MN16 MN17 Genera Asoriculus Mafia Apodemus Orientalomys Centraiomys Mesocricetus

Pseudomeriones Mimomys Dryomimus Vulpes Nyctereules Chasmaporthetes Lynx Machairodus H. longipes-type H. crassum-type Dicerorhinus megarhinus Sus arvernensis Giraffa Apolakki a An o Tourkobounia- 1 Apollonia - 1 Sina p Bayraktep e i l Karaôz û Kalekò y Kayadib i Kemiklitep e Samo s Maramen a Marits a CALT A Megal o Emvolo n Gulyaz i Wolak s Gerakaro u Kemiklitep e D . Dafner o Dytik o Devel i LOCALITIE S Akçako y Metoch i L .0C .8 A Q 1 A+ B D

FIG. 4. — Stratigraphie distribution of some genera and species represented In the Calta fauna.

roots and an occlusal pattern similar to that of stage of evolution of some key taxa, it seems M. dcivakosi. Some Spanish localities yielding this obvious that this fauna should be dated as species were dated by magnetostratigraphy as MN15, and mote probably in the lower part of about 4 Ma old, and the others correlated to this zone. early MN15 (Opdyke et al. 1997; Fejfar et al. In the Tetuel Basin in eastern Spain, Opdyke et 1997). In Spain as well as in othet parts of al. (1997) studied the magnetic stratigraphy of Eutope, late MN15 and later faunas include several sections beating Turolian, Ruscinian and Mimomys species with a more derived dental Villanyian mammal localities. Using magneto- pattern: higher dentine tracts, roots smaller, stratigraphic correlations from the Orrios, increase of the hypsodonty, appearance of Villalba Aha, Escorihuela and Loma del Castillo Dolomys-ipaxttm, etc. This review shows that the sections, they calibrated the MN14/MN15 Calta Mimomys remains among the earliest boundary as 4.18 Ma, and the MN15/MN16 known species of this genus. boundary as correlative to Gilbert/Gauss boun­ Taking into account the bulk of the Calta fauna, dary at 3.58 Ma. They also obtained boundaty and more particularly the time tange and the ages for older (Turolian) and younger

GEODIVERSITAS • 1998 • 20(3) 507 Sen S., Bouvrain G. & Geraads D.

(Villanyian) MN zones which are out of the Bourlière F., Minner E. & Vuattoux R. 1974. — Les scope of the present study. On the other hand, gtands mammifètes de la région de Lamto, Côte d'Ivoire. Mammalia 38 (3) : 433-447. magnetostratigraphy of Pliocene mammal locali­ Bourquin O., Vincent J. & Hitchins P. M. 1971. — ties in Romania calibrated the Dacian/Romanian The vertebrates of the Hluhluwe game reserve. boundary at about 4.2 Ma (Sen 1997); this Corridor (State-Land) Umfolozi game reserve com­ boundary is considered as equivalent to the plex. The Lammergeyer 14: 5-58. MN14/MN15 boundary of the European mam­ Bousquet B. 1992. — Guide des parcs nationaux malian zonation. These results agree in the attri­ d'Afrique : Afrique du Nord et Afrique de l'Ouest. Delachaux & Niestlé (eds), 368 p. bution of an age about 4 Ma to the Calta fauna. Bouvrain G. 1998. — Bovidae, in Sen S. (ed.), Pliocene vertebtate locality of Çalta, Ankata, Acknowledgements Tutkey. 10, Geodiversitas 20 (3) : 467-485. This paper is a result of the field trip organized Btuijn H. de 1976. — Vallesian and Turolian rodents ftom Biotia, Attica and Rhodes (Greece). by É. Heintz in 1972 to which have also partici­ Procedings of the Koninklijke Nederlandse Akademie pated L. Ginsburg and M. Giirbiiz. The sugges­ van Wetenschappen, Amsterdam, B 79: 361-384. tions of the referees J. Michaux and M. Pickford Brunet M., Beauvilain A., Geraads D., Guy F., Kasser greatly improved this paper. M., MacKaye H., Mouchelin G., Sudre J. & Vignaud P. 1997. — Tchad : un nouveau site à Hominidés pliocenes. Comptes rendus de l'Académie des Sciences de Paris, lia, 324: 341-345. REFERENCES Daams R. 1981. —The dental pattern of the dotmice Dryomys, Myomimus, Microdyromys and Peridy- Alcalá Martinez L. 1994. — Macromamiferos neoge- romys. Utrecht Micropaleontological Bulletin, Special nos de la fosa de Alfambra-Teruel. Instituto de Publication, 3: 1-115. Estudios Turolenses. Museo nacional de Ciencias Eisenbetg J. & Lockhart M. 1972. — An ecological naturales, Teruel, 554 p. reconnaissance of Wilpattu National Patk, Ceylon. Anderson J. A., Jermy A. C. & the Earl of Cranbrook Smithsonian Contributions to Zoology 101: 1-118. 1982. — Gunung Mulu national park: a manage­ Eisenmann V. & Sondaat P. 1998. — Hipparions, in ment and development plan. Royal Geographical Sen S. (ed.), Pliocene vertebtate locality of Çalta, Society: 1-345. Ankara, Turkey. 7, Geodiversitas 20 (3) : 409-439. Azanza B., Menéndez E. & Alcalá Martinez L. Feibel C. S., Harris J. M. & Brown F. M. 1991. — 1989. — The Middle-Upper Turolian and Palaeoenvironmental context for the late Neogene Ruscinian Cervidae in Spain. Bolletino della Societa of the Turkana basin: 321-370, in Harris J. M. Paleontológica Italiana 28 (2-3): 171-182. (ed.) Koobi Fora Research Project, volume 3: The Bachelet B. 1990. — Muridae et Arvicolidae Fossil Ungulates: Geology, Fossil Artiodactyls and (Rodentia, Mammalia) du Pliocène du Sud de la Palaeoenvironments. Clarendon Press, Oxford. France: systématique, évolution et biochronologie. Fejfar O., Heinrich W. D., Pevzner M. A. & Thèse de doctorat de l'université de Montpellier II, Vangengeim E. A. 1997. — Late Cenozoic 211p. sequences of mammalian sites in Eurasia: an upda­ Bernick S. 1974. — The community of the wild rumi­ ted correlation. Palaeogeography, Palaeoclimatology, nants in the Gir forest. Ph.D thesis, Yale Univetsity: Palaeoecology, Amstetdam 133: 259-288. 1-226. Fejfar O., Mein P. & Moissenet E. 1990. — Eatly Black C. C, Ktishtalka L. & Solounias L. 1980. — atvicolids ftom the Ruscinian (eatly Pliocene) of Mammalian fossils of Samos and Pikermi. I. The the Teruel Basin, Spain: 133-164, in Fejfar O. & Turolian rodents and insectivores of Samos. Annals Heinrich W. D. (eds), International Symposium of the Carnegie Museum 49: 359-378. Evolution, Phytogeny and biostratigraphy of arvicolids Bocherens H. & Sen S. 1998. — Isotopic investiga­ (Rodentia, Mammalia). Geological Survey, Prague. tion, in Sen S. (ed.), Pliocene vettebrate locality of Geraads D. 1998. — Cervidae et Giraffidae, in Sen S. Çalta, Ankata, Turkey. 11, Geodiversitas 20 (3) : (ed.), Le gisement de vettébtés pliocenes de Çalta, 487-495. Ankara, Turquie. 9, Geodiversitas 20 (3) : 455-465. Bonis L. de, Bouvrain G., Geraads D., Koufos G., Sen Ginsburg L. 1998. — Carnivora, in Sen S. (ed.), Le S. & Tassy P. 1994. — Les gisements de mammi­ gisement de vertébrés pliocenes de Çalta, Ankara, fères du Miocène supérieur de Kemiklitepe, Turquie. 5, Geodiversitas 20 (3) : 379-396. Tutquie. 11- biochronologie, paléoécologie et rela­ Guerin C. & Sen S. 1998. — Rhinocerotidae, in Sen tions paléogéographiques. Bulletin du Muséum S. (ed.), Le gisement de vettébrés pliocenes de national d'Histoire naturelle, Paris, C 16 (1) : Çalta, Ankara, Turquie. 6, Geodiversitas 20 (3) : 225-240. 397-407.

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Submitted for publication on 9 March 1997; accepted on 30 April 1998.

510 GEODIVERSITAS • 1998 • 20 (3) Instructions aux auteurs

cer. Chaque figure doit comportet une échelle Scope of the Journal métrique, sans aucun coefficient multiplicateur Les Geodiversitas publishes papets which concern varied tableaux et gtaphiques, à inclure dans le manuscrit, aspects of Earth Sciences and particularly history of doivent nécessaitement pouvoir êtte imprimés sur sedimentary basins, palaeobiodiversity and paleoen- une page et testet lisibles après réduction éventuel­ vironment. A complete issue of Geodiversitas may le. Des planches en couleut poutront être publiées be devoted to sevetal papers on a single topic undet moyennant une participation finan-cière de ou des the responsability of guest editot(s). Papers with a auteurs. systematic content should follow the International Code of Zoological Nomenclature. Manuscripts, Références bibliographiques without limitation of the numbet of pages, must Denison R. H. 1978. — Placodermi, in Schultze conform strictly with the insttuctions to authots, H. P. (éd.), Handbook of Paleoichthyology, and will be sent to the Editor: Volume 2. Gustav Fischet, Stuttgatt, 128 p. Marshall C. R. 1987. — Lungfish: phylogeny and Service des Publications Scientifiques du Museum, parsimony, in Bemis W. E., Burggren W. W. Geodiversitas, & Kemp N. E. (eds), The Biology and 57 tue Cuviet, Evolution of Lungfishes, Journal of Morphology F-75231 Paris cedex 05 1: 151-162. Tel: (33) 01 40 79 34 38 Schultze H. P. & Atsenault M. 1985. — The pan- Fax : (33) 01 40 79 38 58 detichthyid fish Elpistostege: a close relative to e. mail : [email protected] tetrapods? Paleontology 28: 293-309. Schultze H. P. 1977a. — The origin of the tetra- Instructions to authors pod limb within the rhipidistian fishes: Manuscripts, with illustrations, must be submitted 541-544, in Hecht M. K., Goody P. C. & in ttiplicate (one original and two copies) in A4 Hecht B. C. (eds), Major Patterns in Vertebrate format, double spaced, with margins of at least Evolution. Plenum Ptess, New Yotk and 3 cm and all pages numbered. The original figures London. should be sent with the revised manuscript, as well as a 3.5" diskette Apple Macintosh ot IBM-compa­ Epreuves et tirés à part tible (Word, Wotd Petfect...) format, which will Les épteuves seront adressées à l'auteur ou au pre­ also contain tables and possibly figures (Adobe mier auteur (sauf indication conttaite) et devront Illustratot, Photoshop; Deneba Canvas). être retournées cottigées sous huitaine. Les correc- tions, auttes que celles imputables à la rédaction ou Format à l'imprimeur, seront à la charge des auteurs. Le(s) Papets are to be written in simple and concise auteur(s) recevront gracieusement vingt-cinq tirés à French ot English. They should be otganized as fol­ part, les tités à part supplémentaires seront à com­ lows: mander sur un formulaire joint aux épteuves. - a brief title in French and English; - a suggested tunning head; Soumettre un atticle pour publication dans - name(s) of author(s), followed by their full pro­ Geodiversitas suppose que celui-ci ou tout article fessional address(es) and, if possible, Fax number proche dans la même langue ou une autre langue, and e-mail; n'ait pas été soumis dans une autre revue, même - absttacts (in English and French) not exceeding dans l'attente de son acceptation. Les droits de 800 signs each, with key wotds and "mots cles"; reproduction de l'atticle, y comptis des illustra­ - text with italicized wotds for Latin: taxa of gene­ tions, sont téservés à la revue. La reproduction de ric and specific ranks (e.g. Cellaria Ellis et Solander, tout ou partie de l'article doit faire l'objet d'une 1786) etal; demande écrite ptéalable adressée à la rédaction. - references to authors in main text should be pre­ sented, in lower case, as follows: Smith (2001), Smith (2001, 2002), (Smith 2001), (Smith 2001; Cary 2002), (Smith & Caty 2003, 2005), Smith (2001: 1), Smith (2001, fig. 2); - references to illustrations and tables should be indicated as follows: (Fig. 1), (Fig. 2A, D), (Fig. 2A-C), (Figs 3, 6), (Figs 3-5), (Table 1);

512 GEODIVERSITAS • 1998 • 20(3) Instructions aux auteurs

— keep acknowledgements short and place them at Marshall C. R. 1987. — Lungfish: phylogeny and the end of the text before references; please do not parsimony, in Bemis W. E., Burggren W. W. forget the tevisers; & Kemp N. E. (eds), The Biology and Evolution of Lungfishes, Journal of Morphology - give captions to illusttations on a separate sheet. 1: 151-162. Schultze H. P. & Arsenault M. 1985. — The pan- Illustrations derichthyid fish Elpistostege: a close telative to The editotial board will pay special attention to the tettapods? Paleontology 28: 293-309. quality and televance of illustration. Schultze H. P. 1977a. — The otigin of the tetra- Line dtawings must be in Indian ink ot high quali­ pod limb within the thipidistian fishes: ty laset ptintouts; high contrast photogtaphs, pla­ 541-544, in Hecht M. K., Goody P. C. & ced on white or black backgrounds, are required. Hecht B. C. (eds), Major Patterns in Vertebrate These can be grouped into figures and identified Evolution. Plenum Ptess, New York and London. by letters A, B, C ... Plates ate not placed at the end of the article: they will be considered as figures Proofs and reprints and numbered as such. Arrange figures to fit one ot Proofs will be sent to the first authot for correction two columns (70 x 190 mm ot 144 x 190 mm). and must be tetutned within eight days by express Associate interpretation of photogtaph with line mail. Authors will receive twenty-five offprints free dtawing. No diagram or table is to exceed one of charge; furthet offprints can be ordered on a page. Lettets, numbers, etc., for each figure, are to form supplied with the proofs. be indicated on an accompanying ovetlay, not on the original figure (line cut or half-tone). A scale The submission of a manuscript to Geodiversitas bar is needed for each figure (without magnifica­ implies that the papet, ot a similar one, is not being tion factor). offeted for publication elsewhete. Copyright of References published papet, including the illusttations, Denison R. H. 1978. — Placodermi, in Schultze becomes the property of the journal. Requests to H. P. (ed.), Handbook of Paleo ichthyology, teptoduce material from Geodiversitas should be Volume 2. Gustav Fischer, Stuttgart, 128 p. addressed to the editor.

GEODIVERSITAS • 1998 • 20(3) 513 Mise en page Noémie de la Selle Packaging Éditorial

Achevé d'imprimer sur les presses de l'Imprimerie Durand 28600 Luisant (France) Dépôt légal n° 10259

Printed on acid-free paper Imprimé sur papier non acide

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geodiversitas 1998 • 20 (3)

Pliocene vertebrate locality of Calta, Ankara, Turkey Edited by Sevket Sen

,„ — Lunkka J. P., Kappelman J., Ekart D. & Sen S. Pliocene vertebrate locality of Calta, Ankara, Turkey. 1. Sedimentation and lithostratigraphy

— Janoo A. & Sen S. Pliocene vertebrate locality of Calta, Ankara, Turkey. 2. Aves: Struthionidae

ÎSÎ • Reumer I- w- F- Pliocene vertebrate locality of Çalta, Ankara, Turkey. 3. Insectivores

Sen S. 359 • Pliocene vertebrate locality of Calta, Ankara, Turkey. 4. Rodentia and Lagomorpha

379 • Ginsbur9 L Le gisement de vertébrés pliocenes de Çalta, Ankara, Turquie. 5. Carnivores

Guérin C. & Sen S. 397 • Le gisement de vertébrés pliocenes de Çalta, Ankara, Turquie. 6. Rhinocerotidae

Eisenmann V. & Sondaar P. 409 •

Pliocene vertebrate locality of Çalta, Ankara, Turkey. 7. Hipparion

441 • Guérin C., Faure M. & Sen S.

455 • Le gisement de vertébrés pliocenes de Çalta, Ankara, Turquie. 8. Suidae

Geraads D. 467 •

Le gisement de vertébrés pliocenes de Çalta, Ankara, Turquie. 9. Cervidae et Giraffidae

487 • Bouvrain G.

Sen S., Bouvrain G., Geraads D. Le gisement de vertébrés pliocenes de Çalta, Ankara, Turquie. 10. Bovidae 497 • Pliocene vertebrate locality of Calta, Ankara, Turkey. 12. Paleoecology, biogeography and biochronology

Bocherens H. & Sen S. Pliocene vertebrate locality of Çalta, Ankara, Turkey. 11. Isotopie investigation

Conception Graphique : Isabel Gautray ISSN : 1280-9659

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