Miocene vegetation and climate dynamics in Eastern and Central Paratethys (Southeastern Europe)

Palaeogeography, Palaeoclimatology, Palaeoecology 304 (2011) 262–275

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Palaeogeography, Palaeoclimatology, Palaeoecology

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Miocene vegetation and climate dynamics in Eastern and Central Paratethys(Southeastern Europe)

Dimiter Ivanov a,⁎, Torsten Utescher b, Volker Mosbrugger c, Svetlana Syabryaj d,Desa Djordjević-Milutinović e, Serge Molchanoff d

a Institute of Botany, Bulgarian Academy of Sciences, Acad. G. Bonchev Str. 23, BG-1113 Sofia, Bulgariab Steinmann Institute, Bonn University, Nußallee 8, D-53115 Bonn, Germanyc Senckenberg Research Institute and Natural History Museum, Senckenberganlage 25, D-60325 Frankfurt, Germanyd Institute of Geological Sciences NAS of Ukraine, O. Gonchara str. 55-b, Kiev 01601, Ukrainee Natural History Museum, Njegoseva 51, 11000 Belgrade, Serbia

⁎ Corresponding author. Tel.: +359 29793773; fax: +E-mail address: [email protected] (D. Ivanov).

0031-0182/$ – see front matter © 2010 Elsevier B.V. Adoi:10.1016/j.palaeo.2010.07.006

a b s t r a c t

Article history:Received 16 November 2009Received in revised form 3 July 2010Accepted 9 July 2010Available online 15 July 2010

Keywords:VegetationClimatePalynologyPalaeobotanyCoexistence approachMioceneParatethysSoutheastern Europe

From the Cretaceous to modern time the earth's climate system has changed repeatedly from a greenhouseto an icehouse. The Miocene appeared as the last warm episode in earth history, prior to the establishment oflarge Northern Hemisphere ice sheets. Furthermore, the Miocene was characterized by both, large-scalemarine transgressions and pronounced episodes of regressions, e.g. opening and closure of marine corridorsas well as appearance and disappearance of lakes and swamps. The sea-level changes and instability of theclimate system strongly influenced not only marine ecosystems but also the terrestrial biota.To throw new light on the Miocene evolution of European ecosystems and climate dynamics, we comparedplant assemblages from southeastern Europe. Palaeogeographically this area is a transitional belt betweenTethyan and Paratethyan realms, and it thus constitutes a crucial geographical position when one tries tocomprehend the exchange pathways of various floristic elements, appearance and evolution of steppevegetation, interaction between forest and steppe vegetation, and of course the corresponding climatesinfluencing the vegetation dynamics.Both macro- and microfloristic plant associations were considered in our analysis. Large palaeofloristic andpalaeocoenotic transformations occurred during the Miocene. The Early to Middle Miocene flora is rich anddiverse in thermophilous elements, which formed polydominant mesophytic to hygromesophytic forests.Early Miocene climate was warm and humid with mean annual temperatures mainly above 16 °C and annualrainfalls over 1000 mm.The Middle Miocene was the warmest period of the whole Miocene, with annual temperatures ranging from17 °C to19 °C andwinter temperatures from7 °C to 12.5 °C. Climatic changes after theMiocene climatic optimumcaused changes in floristic composition and vegetation structure. The vegetation shows a decreasing trend inabundance of palaeotropic and thermophilous elements, reduction of macrothermic elements, and disappear-ance of evergreen laurel forests. Together with these changes is a corresponding increase in the role ofarctotertiary species in plant communities, and they became dominants in mesophytic forests.The available data indicate that major vegetation changes occur in the late Miocene. This period is characterizedbymore diverse climatic conditions,whichweredirectedbyglobal climatic changes andprobably complicatedbyregional palaeogeographic reorganizations and tectonic processes. Slight cooling and some drying is recordedfor the beginning of the Late Miocene, followed by fluctuations of palaeoclimate parameters observed whichdisplay cycling change of humid/dryer and warmer/cooler conditions. Our study provides a new insights on theMiocene vegetation and climate dynamics and evolution in Southeastern Europe.

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1. Introduction

The Cenozoic is a time in plant evolution in Eurasia whenPalaeogene vegetation dominated by evergreen and thermophilouselements was replaced by vegetation dominated by deciduous and

temperate plants. The processes of plant evolution and transformationof vegetation and coenotic structure was significantly forced bychanges in the earth climate system and the transition from agreenhouse to icehouse world. In this complicated and intriguingevolutionary process the Miocene period appeared as the last warmepisode in earth history, prior to the establishment of a large NorthernHemisphere ice sheet. In fact, the Miocene is considered the mostcritical interval in the build-up of ice masses on land (Zachos et al.,

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2001). Furthermore, the Miocene was characterized by both, large-scale transgressions and pronounced regressions, e.g. opening andclosure of marine corridors, appearance and disappearance of lakesand swamps in Europe (Kojumdgieva and Popov, 1989; Meulenkampet al., 1996; Rögl, 1998, 1999, 2001; Meulenkamp and Sissingh, 2003;Ilyina et al., 2004a,b; Harzhauser and Piller, 2007; Harzhauser et al.,2007; Harzhauser and Mandic, 2008). Alpine tectonics was veryactive, forcing the uplift of the Carpathians, Dinarides, Balkans,Rhodopes, and Caucasus. The collision of the Afro-Arabian andEurasian plates resulted in closure of the ancient Tethys Sea (Allenand Armstrong, 2008, and references therein), which was just oneaspect in the plate reorganization and oceanographic changes. Thiscollision caused the closure of the Tethys Gateway that previouslylinked the Atlantic and Indian Oceans. The sea-level changes, tectonicactivities, plate reorganizations, changes in ocean circulations, andinstability of the climate system strongly influenced both, marineecosystems and the terrestrial biota, and they collaborated to developdistribution of vegetation patterns seen today.

To throw new light on the Miocene evolution of Europeanecosystems and climate dynamics, we compared plant assemblagesfrom southeastern Europe. The geographical position of this area,located between Africa and Eurasia and between a Mediterraneanand continental temperate climate, makes it of great interest forpalaeobotanical studies. It is also a key area to comprehend theexchange pathways of various floristic elements, appearance andevolution of steppe vegetation, interaction between forest andsteppe vegetation, and of course the corresponding climatesinfluencing the vegetation dynamics. Several studies were carriedout in the area of the Balkans and southeastern Europe, aiming toelucidate the vegetation and climate evolution. Most studies arebased mainly on macrofloristic or microfloristic records, but in rarecases they combine all available data. In analysis of vegetation andclimate, studies on individual floras or on local groups of florasprevail (Ivanov et al., 2002; Kovar-Eder et al., 2006; Ivanov et al.,2007a,b; Syabryaj et al., 2007; Utescher et al., 2007a; Ivanov et al.,2008; Kovar-Eder et al., 2008). However, comparative, regionalstudies are also available (Stuchlik et al., 1999; Bruch et al., 2004,2006; Fauquette et al., 2007; Jiménez-Moreno and Suc, 2007;Jimenez-Moreno et al., 2007), which in most cases deal withlatitudinal climatic and vegetational changes or consider distincttime intervals. But for the Miocene, besides a latitudinal vegetation/climate differentiation, a change in vegetation patterns in an east–west direction was observed (e.g. see in Mai, 1989, 1991, 1995;Kovar-Eder et al., 1996; Utescher et al., 2007b).

In the present study we analyze and compare vegetational andclimatic changes throughout the Miocene along the seacoast ofParatethys, taking into consideration the available palaeobotanicaldata from Ukraine, Bulgaria, and Serbia (Fig. 3). These three countrieshave a key geographic position in southeastern Europe. Ukraine linkscentral and eastern Paratethys during the Miocene. Its western partbelongs to the Ukrainian Carpathians (Central Paratethys), while theUkrainian plain and Crimea belong to the eastern Paratethys. Thus aneast–west land bridge existed in Ukraine connecting Central Asia toCentral Europe.

Bulgaria lies in the transitional area between the Tethyan andParatethyan realms and constitutes a crucial geographical position inattempts to comprehend the evolution of the recent Mediterraneanvegetation and the exchange paths of various floristic elementsbetween northern and southern Europe that provide a bridgebetween Asia Minor and Europe, enabling the appearance andevolution of grasses and steppe vegetation in Europe under favourableclimatic conditions. And finally, the specific palaeogeographic positionand geodynamic evolution of Serbia makes it a major migration routeof plant species from the Balkans to the Pannonian lake system and aregion of exchange of plant diversity between central and southeast-ern Europe along the Dinarides mountain chain.

Today the eastern part of the study area is mainly occupied bysteppe, while the more southerly regions (e.g. along the Black Seacoast) are refuge areas of some thermophilous plants. However, whatwas the vegetation in this region before Northern Hemisphereglaciations, especially at the time when important palaeogeographicchanges in the Middle to Late Miocene led to the evolution ofvegetation patterns and vegetation distribution that are different fromthose of today? It is important to outline the scenario of Miocenedevelopment and regional differentiation of the southeastern Euro-pean vegetation that was the result of non-linear climatic coolingforced by palaeogeographic change.

2. Geology and palaeogeography of eastern andsoutheastern Europe

The palaeofloras analyzed are geographically distributed in marineand fresh water basins belonging to the Central and EasternParatethys (Fig. 1). The appearance of the Paratethys in the beginningof the Oligocene was strongly affected by the collision of the Afro-Arabian and Eurasian plates and by Alpine tectonics (Allen andArmstrong, 2008). Initially it developed as an epicontinental seaoccupying territories between 40 and50° north latitude, and it spreadfrom the Rhone Basin in France in the west to Central Asia in the east.During the Neogene the Paratethys displayed a long-term trend ofdecreasing marine influence and a correlative reduction in size of themarine depositional domains (Meulenkamp and Sissingh, 2003). Atthe end of the Early Miocene it was segregated into threepaleogeographic and geotectonic units (sub-basins), namelyWestern,Central and Eastern Paratethys. The Western Paratethys was thesmallest and comprises the Alpine Foreland Basins. The CentralParatethys includes Pannonian Basin and Carpathian Foreland, andthe Eastern Paratethys comprises the Euxinian Basin (Black Sea),Caspian Sea, and Aral Sea (Piller et al., 2007). Part of the CarpathianForeland changed its geo- and hydrodynamic regime at the end of theMiddle Miocene, switching from the central Paratethys into theeastern Paratethys domain. The palaeogeographic dynamics in thisarea include large-scale transgressional/regressional cycles, openingand closure of marine corridors, appearance and disappearance oflakes and swamps (Kojumdgieva, 1983; Kojumdgieva and Popov,1989; Rögl, 1998; Popov, 2001; Meulenkamp and Sissingh, 2003;Goncharova et al., 2004; Ilyina et al., 2004a,b; Harzhauser and Piller,2007; Harzhauser and Mandic, 2008).

Due to the changing seaways or separations between the Caspian,Euxinian, Forecarpathian (Dacian) and Pannonian Basins throughoutthe Neogene (Rögl, 1998) facies and fossil content of the sedimentswere quite different. Hence, the independent evolution of thedifferent sub-domains of the Paratethys led the construction ofseveral regional stratigraphic schemes. These schemes are largelybased on the mollusks, ostracods, and benthic and planktonicForaminifera, reflecting the endemic character of the fauna in separatebasins. Reliable correlations are established between the Central andEastern Paratethys regional stratigraphies and the Mediterraneanstandard stratigraphy (Fig. 2) according to magnetostratigraphic data,isotopic dating, nannoplankton evidences etc. (e.g. Vasiliev et al.,2004, 2005; Popov et al., 2006; Snel et al., 2006a,b; Vasiliev, 2006;Harzhauser and Piller, 2007; Kovac et al., 2007; Piller et al., 2007). Asmentioned above, the study area belongs to two palaeogeographicdomains — (1) the eastern part to the Central Paratethys (Transcar-pathian, Carpathian Foredeep and Forecarpathian Basin, and thesouthern part of Pannonian Basin System), and (2) the EasternParatethys (Ukrainian plane, Crimea and NE Bulgaria).

Shallow-water sedimentation existed in the Carpathian Foredeepduring the Egerian stage in the area of the Ukraine Carpathians(Central Paratethys, Precarpathian/Forecarpathian Basin). At thattime lagoonal and continental deposits accumulated. Later duringthe Eggenburgian a sea transgression restored marine conditions in

Fig. 1. Palaeogeographic sketch-maps of the Mediterranean and Paratethyan area (afterRögl, 1998; Popov et al., 2004; Harzhauser and Piller, 2007): a) Early Miocene; b)Middle Miocene; c) Late Miocene. The dotted line marks approximate outlines ofstudied area.

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the area. Alpine orogenesis stimulated the continuing rising of theCarpathian and Crimea mountains and the subsidence of the foothillsadjacent to the platform margins at the beginning of Middle Miocene.This changed the land-sea distribution in the Central and EasternParatethys areas profoundly, and during the Middle Miocene the seatransgression took place. This was the time when the ForecarpathianBasin had a maximum extension, and to the south it covered theterritory of northwestern Bulgaria (the so-called Dacian Gulf ofForecarpathian Basin) ( Kojumdgieva and Popov, 1989; Rögl, 1998;Ivanov et al., 2002). At that time the extension of the basin continued

to the west through the territory of Serbia, thus providing marineconnection with the Vienna-Pannonian Basin. Regression and isola-tion of the basin took place at the end of the Badenian stage, caused bythe uplift of the Carpathian Mts.

Early Volhynian transgression occurred in the southern part of theForecarpathian Basin (NWBulgaria) and in the Precarpathian Foredeep.The connection to the Pannonian Basin was closed, and from theVolhynian onwards until the end of the Miocene a seaway to theeastern Paratethys developed (Rögl, 1998). After a regressive during themiddle and upper Chersonian, the sea returned in the early Maeotian,and the basin occupied the southern part of the Forecarpathian Basinuntil the Upper Pontian and Lower Pliocene, with gradual replacementof marine/brackish sedimentation by freshwater and deposition ofcontinental sediments and coals (Ivanov et al., 2002). The Transcar-pathian Basin also became shallow during the Pannonian-Pontian, andit was occupied by fresh water.

In the Eastern Paratethys, tectonic movements at the Oligocene/Miocene boundary caused uplift and sea regression in the EuxinianBasin— e.g. Ukrainian plain, Crimea Peninsula, and the Varna–Balchikdepression (Ivanov et al., 2007c; Syabryaj et al., 2007).

A tectonic reorganization in the latest Early to earliest MiddleMiocene in the eastern Paratethys resulted in the emergence of theGreater Caucasian archipelago (Meulenkamp and Sissingh, 2003),followed by a transgression north and west of the Black Sea. The firsttransgression took place in the Tarkhanian, followed be a regressivetrend in the Tshokrakian. During that time the Ponto-Caspian areawas sealed off again from the open oceans (Rögl, 2001), thus leadingto the appearance of brackish environments.

During the early Konkian full marine conditions returned in theEuxinian basin with stenohaline molluscs and foraminifers, when theIndian Ocean was linked to the Paratethys in eastern Anatolia (Rögl,1998, 2001). The Euxinian Sea transgressed onto large land areasduring the Volhynian–Chersonian, especially at the northern andeastern margins (Paramonova et al., 2004). During the Chersonian aminor regression occurred, and before the end of this stage the seaabandoned northeastern Bulgaria, but a new transgression took placeto the north (Ukraine), and sedimentation continued in the Maeotianuntil the Pontian, when a reduction of the marine basin took place. Atthe end of the Pontian the outlines of the basin were close to present.

From a palaeogeographical point of view the Serbian area belongsto two domains — Central and Eastern Paratethys. The easternParatethys occupied only a small area in eastern Serbia. The mostimportant structure was the Dinarid Lake System formed during theLate Oligocene and Miocene (Krstić et al., 2003; Harzhauser andMandic, 2008) — intramountain basins parallel to the Dinaridmountain chains developed in NW–SE direction. Subsequent riftingin the Pannonian Basin System triggered themarine flooding of part ofthe Dinarid Lakes and considerably reduced them. After thedisintegration of the Paratethys Sea at about 11.6 Ma and develop-ment of the Pannonian lake the northern part of Serbia became a partof the Pannonian Basin. At the end of the Miocene only relict lakesexisted in northern Serbia (e.g. Paludina Lake) as a residual from thesouthernmost part of the Pannonian lake system.

3. Materials and methods

The material from Ukraine analyzed in this study comes fromMiocene sediments of two regions (Fig. 3) in the northeastern part ofthe Central Paratethys (Carpathian Foredeep), and the northern partof the Eastern Paratethys (Ukraine Plain and Crimea). Regional stagesas well as stratigraphic units for each of the above regions areshown in Fig. 2. Palynological data come from complete stratigraphicsequence of the Miocene deposits (Shchekina, 1979; Syabryaj andShchekina, 1983). The samples analyzed were collected from cores ofnumerous wells as well as from outcrops (Fig. 3).

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The Early Miocene flora of Bulgaria is not sufficiently well studied.At that time the territory of the country was an upland area, and nosediments are known. Only in the area of the Bobovdol Basin (near thevillage of Babino) did a fresh water lake exist with a fossil flora datingfrom the Late Oligocene–Early Miocene transition (equivalent to theEgerian flora in the Central Paratethys (Palamarev et al., 1998). TheMiddle Miocene palaeoflora is studied from several localities:Forecarpathian and Euxinian Basins (Paratethyan realm) and freshwater basins of Chukurovo, Satovcha, Slasten, and Simitli. The dataabout the Late Miocene flora and vegetation come from theForecarpathian Basin (brackish environment) and from several freshwater localities, e.g. Sandanski, Gotse-Delchev, Tundzha, Staniantsi,Beli breg, Sofia, Karlovo, Kyustendil, and Razlog Basins (Fig. 3). Bothmacro-and micropalaeobotanical studies have been undertaken inthese basins.

The analysis and discussion of the vegetation and climate in Serbiawas carried out on the already published lists of macroflora fromnumerous localities (see Table 1.). These lists and the additionalcheckup of already published materials were used to determine themain floristic elements and reconstruct vegetation dynamics. Most ofthe material consists of leaf imprints, but there were also some fruit/seed, and permineralized branches and conifer cones. Palynologicaldata have also been used in floristic/climate reconstructions, but onlyfor the Pontian vegetation from the southern coast of the PannonianBasin. The detailed overview of vegetation replacements and climatefluctuations during the Miocene of Serbia was presented in Utescheret al. (2007a).

To study the palaeoclimate evolution in different parts of theParatethian realm during the Miocene the Coexistence Approach (CA)is used. The method is described in detail in Mosbrugger and Utescher(1997). The CA follows the nearest living relative (NLR) concept. Thedistribution of plant species depends strongly on climatic conditions.The climatic tolerances of fossil plants is considered to be close to theirNLR's. This allows the use of transfer functions to estimate pastclimates based on the fossil plant record. By using the database ofextant taxa and their climatic requirements (PALAEOFLORA: Utescherand Mosbrugger, 1990–2007), “coexistence intervals” of differentclimatic parameters could be calculated that allow the majority ofencountered fossil taxa to exist at that location. The overlap in theranges of the different taxa defines the interval of coexistence, whichis then interpreted as the range of the palaeoclimate. The reliability ofthe results increases with the percentage of taxa that may coexist inthis interval (Mosbrugger and Utescher, 1997). In the present studythe mean annual temperature (MAT), cold month mean temperature(CMMT), warm month mean temperature (WMMT), mean annualprecipitation (MAP), precipitation in the warmest month (MPwarm),precipitation in the driest month (MPdry), and precipitation in thewettest month (MPwet) were analyzed and discussed.

4. Results and discussion

4.1. Vegetation

4.1.1. The Early Miocene

4.1.1.1. Ukraine. Palynological complexes from the Lower Miocenedeposits are not uniform. The vegetation record of the Eggenburgianindicates a significant warming during the transgressive cycle, whenin the lowlands diverse broadleaved mesophytic forests with

Fig. 2. Regional stages of the Central and Eastern Paratethys (according to Gradsteinet al., 2004a; Gradstein et al., 2004b; Ogg et al., 2008) with stratigraphic subdivision forthe Eastern and Central Paratethys (Harzhauser and Piller, 2007; Harzhauser et al.,2007; Piller et al., 2007; Zuschin et al., 2007; Harzhauser andMandic, 2008; Harzhauseret al., 2008).

Fig. 3. Location of fossil floras in the studied area: 1. Ukraine; 2. Bulgaria; 3. Serbia.

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evergreen species existed. Magnolia, Lauraceae, Engelhardia, Ilex,Aralia, Vitis, Symplocaceae, Staphyleaceae, Rosaceae, Sapotaceae,Arecaceae, Myrtaceae, Theaceae, Gleicheniaceae and Schizaeaceaewere the main components. Swamp forests occupied only limitedareas. The spore and pollen record of both, the Precarpathian andTranscarpathian areas indicates the presence of altitudinal vegetationzones. Carpathian Mts are supposed to have been a low mountainchain at the beginning of the Early Miocene (Egerian), probably notmore than 1000 m in altitude, in accordance with this assumption.The uplift of the Carpathians forced the vertical differentiation of plantcommunities, becoming more pronounced at the end of the Egerian(Syabryaj and Stuchlik, 1994; Syabryaj et al., 2007). Temperate plantsexisted in the mid-elevation and partially in the upper altitudinalvegetation belts. Picea, Abies, Tsuga, and Podocarpus were mostimportant in the coniferous belt covering the mountain slopes.Warm-temperate representatives of Juglans, Acer, Tilia, and Quercuswere abundant in broadleaved forests. As a whole the vegetation datasuggest a warm and humid climate for the Transcarpathian area ofUkraine during the Eggenburgian (Syabryaj, 1981).

Significant differences existed in the composition of the flora andvegetation from the mountain slopes and plains. The main vegetationunits growing in the plains were subtropical mesophytic forests andswamp communities with abundant Myricaceae. The undergrowth inthese forests was composed of subtropical ferns of the Schizaeaceae,Gleicheniaceae, Cyatheaceae, Matoniaceae, and Dicksoniaceae fami-lies. Mixed coniferous and broadleaved forests with thermophilousspecies of Juglandaceae and Fagaceae (Castanea) were most commonon the inner slopes of the Carpathians (Syabryaj, 1980).

In the regressive cycle of the Ottnangian, no plant remains havebeen found. A poor spore and pollen complex was recovered for thelate Carpathian, indicating the presence of broadleaved forest withFagus and Liquidambar. Meanwhile a reduction of thermophilousfloristic elements in plant communities occurred (Syabryaj, 1991).

The coastal areas in the south of Ukraine were covered by swampforests with Taxodium, Nyssa, and Myrica in the Sakaraulian(=Eggenburgian/Ottnangian). Riparian vegetation occupied rivervalleys. It was composed by broadleaved forests with abundantJuglandaceae (Juglans, Pterocarya, and Platycarya) and some repre-sentatives of the Betulaceae family (Carpinus and Alnus). The presenceof Engelhardia, Vitaceae, Theaceae, and Myrtaceae is evidence for awarm and moist climate, similar to that in the Carpathian region.Polydominant forests of Quercus, Moraceae, Tilia, Betula, Fagus,Castanea, Ulmus, Acer, Magnolia, Sapindus, Fabaceae, and Rosaceae

spread over continental areas (Syabryaj et al., 2007). Ferns decreasedin the fossil record, and terrestrial herbs became more important.The sea regression at the end of the early Miocene (Kotsakhurian =Carpathian= end of Burdigalian) forced the expansion of halophilous(Chenopodiaceae) vegetation on saline soils. At the same time itcaused the reduction of Taxodiaceae swamp forest. Changes in pollencontent indicate some seasonality of precipitation (for details seeShchekina, 1979).

4.1.1.2. Bulgaria. During the early Miocene the area of Bulgaria wasupland, and no sediments are known. Only in the area of BobovdolBasin (near the village of Babino) did a smaller fresh water lake existwith a fossil flora dating to the Late Oligocene–Early Miocenetransition (equivalent to the Egerian flora in the Central Paratethys(Palamarev et al., 1998). Swamp forest with Glyptostrobus, Taxodium,Comptonia, Myrica, Cyrilla, Andromeda, and Nyssa existed in floodedareas surrounding palaeolakes. Hygromesophytic to mesophyticforest and shrub palaeocoenoses composed of representatives ofLauraceae, Fagaceae, Juglandaceae, Mastixiaceae, Betulaceae, Tiliaceaeetc. were widespread. An increased abundance of thermophilous taxabelonging to the Lauraceae, mass occurrences of Sideroxylon andEngelhardia, and a diminished importance of arctotertiary elements(e.g. disappearance of representatives of the genera Acer, Alnus,Carpinus, and Populus) indicate warming at the beginning of the earlyMiocene (Bozukov et al., 2009).

4.1.1.3. Serbia. Floras of the Early Miocene of Serbia have alwaysincluded both the evergreen broadleaved and the deciduous compo-nents, with a very common addition of xerophilous elements. The bestrepresented subtropical taxa were Lauraceae: Laurus, Ocotea, Persea,and Daphnogene (sporadically). The evergreen component is domi-nant in most floras. Although at first sight these seem to be typicalsubtropical associations, deciduous elements are almost alwayspresent to various extents. These are mostly the species characteristicof riparian forests: Alnus, Salix, Populus, and probably some species ofMyrica. Besides the usual riparian species, the deciduous elementsmost commonly include Acer, Castanea, Zelkova, Ziziphus, and variousJuglandaceae. The deciduous species typical of colder areas or higheraltitudes, such as Fagus and Betula, are recorded rarely.

According to this composition of paleofloras it could be concludedthat the zonal vegetation during most of early Miocene was repre-sented by mixed mesophytic forests rich in evergreen components.The (semi-) xerophyte elements are represented by Leguminosae

Table 1Localities with fossil flora (macroflora and pollen) used in vegetation reconstructions andpalaeoclimate interpretations [according to: Shchekina, 1979; Syabryaj and Shchekina,1983; Syabryaj et al., 1993, 2007; Syabryaj and Stuchlik, 1994 (Ukraine); Palamarev, 1982,1989, 1990, 1994a,b; Bozukov, 1996, 2001, 2002; Bozukov and Palamarev, 1995; Ivanov,1995, 2003; Palamarev and Ivanov, 1998, 2004; Palamarev et al., 1998, 1999, 2002;Stuchlik et al., 1999; Ivanov et al., 2002, 2007a,b,c, 2008; Ivanov and Slavomirova, 2004;Palamarev and Bozukov, 2004; Bozukov et al., 2009; Utescher et al., 2009a (Bulgaria);Pantić, 1956, 1957, 1989; Pantić and Dulić, 1992a, b, 1993; Pantić and Mihajlović, 1976 ;Mihajlović, 1988, 1991;Mihajlović and Lazarević, 1995, 1999, 2003; Utescher et al., 2007a,b; Djordjević-Milutinović and Ćulafić, 2008 (Serbia)].

Country/Region

Locality Age

UkraineUkraine Plane10 Chaplinka Pontian9 Emetovka Bessarabian; Early Maeotian8 Korobki Tshokrakian; Karaganian; Konkian7 Leniskoye, Crimea Tarkhanian6 Bolshi Kopani Karadzhalganian; SakaraulianUkrainian Carpathians5 Velikaya Began Pannonian, Pontian4 Beregovo Middle Sarmatian3 Ternova Late Badenian3 Olkhovtsy Early Badenian2 Voditsa Eggenburgian1 Zaluzh Egerian

Bulgaria18 Razlog Late Miocene (Pontian, Pliocene?)17 Sofia Late Miocene ( Pontian, Pliocene)16 Beli breg Late Miocene (Pontian, Pliocene?)15 Staniantsi Late Miocene (Latest Maeotian, Pontian)14 Tundzha Late Miocene (Late Miocene: Pontian)13 Gotse-Delchev Late Miocene (Pontian, Pliocene?)12 Kyustendil

(Nikolichevtsi)Late Miocene (Maeotian)

11 Sandanski Basin Maeotian10 Drenovets Bessarabian9 Koshava Bessarabian8 Ruzhintsi Volhynian7 Simitli Middle Miocene (Badenian)6 Slasten Middle Miocene (Badenian)5 Satovcha Middle Miocene (Badenian)4 Chukurovo M5iddle Miocene (Badenian = Langhian)3 Forecarpathian Basin

(well)Badenian, Volhynian, Bessarabian, Chersonian,Maeotian, and Early Pontian

2 Euxinian Basin(well)

Oligocene; Tarkhanian, Karaganian, Bessarabian,and Chersonian

1 Bobovdol-Babino Latest Oligocene–Early Miocene

Serbia21 Ćirkovac (Kostolac) Pontian20 Kolubara (Field D) Pontian19 Kolubara (coal mine) Pontian18 Dubona II Late Pannonian17 Sremska Kamenica Early Pannonian16 Pančevo bridge

(Beograd)Sarmatian

15 Bela Stena (Valjevo) Early Sarmatian14 Boždarevac Late Badenian to Early Sarmatian13 Bukovac (Saranovo) Late Badenian to Early Sarmatian12 Selište Badenian to Early Sarmatian11 Lešće (Beograd) Late Badenian10 Misaca Badenian (Upper part)9 Slanci Badenian?8 Melnica Badenian7 Popovac Badenian (Lower part)6 Zlatokop Early Miocene5 Kaludra Early Miocene4 Snegotin-Vukovići Early Miocene (Burdigalian)3 Stamnica Eggenburgian, Ottnangian2 Ćićevac Latest Chattian1 Ravna Reka Latest Chattian

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and mediterranean-type Quercus. The dominant conifers are Sequoiaand Pinus, while the typical swamp conifers Taxodium and Glyptos-trobus are poorly represented. Changes in the palaeogeographic

situation influenced the floristic composition, mainly by the evolvingpalaeo-relief and difference in palaeo-altitude. In the Early Miocene,the relief of the northern part of Serbia was only slightly increased,while in the southern part, in the area of the present-day DinaridMountains, it was moderately raised, but no high mountains existed(Pantić and Dulić, 1992a).

4.1.2. The Middle Miocene

4.1.2.1. Ukraine. Broadleaved forests with Ulmus, Castanea, andfrequent Engelhardia dominated the Middle Miocene vegetation inthe Ukrainian Carpathians. The existence of a seaway between theCentral Paratethys and the World Ocean during the Badenian and theappearance of warm oceanic currents favoured the distribution ofthermophilous terrestrial plants. Nevertheless, a slight trend to adecrease in the number of warm-loving plants (except Engelhardia)towards the end of Middle Miocene is evident. Picea and Ulmusbecame more important in the mountain forests.

During the late Badenian, humid coastal and riparian forests withCarya, Pterocarya, Liquidambar, Alangium, and different species ofMyrica, Comptonia, and Daphnogene (leaf remains) played animportant role in the Ukrainian Carpathians (Iljinskaya, 1960,Shvarieva, 1983). The mixed forests included Juglans, Engelhardia,Tilia, Acer, Castanea, Fagus, Quercus, and Platycarya, as well as Zelkovaand Ulmus. The shrub layer became more diverse (Lonicera, Diervilla,Elaeagnus, Corylus, and Cornaceae). Increasing percentages of herba-ceous pollen is interpreted by Syabryaj et al. (2007) as expansion ofmountain meadow communities in the mountainous areas beinguplifted at that time.

The beginning of the Sarmatian was characterized by intensiveorogenic movements, volcanism, and a reduced extension of themarine basins in the Ukrainian Carpathians. In general, the broad-leaved deciduous forests were floristically similar to those of the lateBadenian, but beech communities with oak–beech and beech–chestnut assemblages appeared. The species of Lauraceae (Daphno-gene) and some subtropical ferns (e.g. Gleicheniaceae) were maincomponents of the undergrowth in broadleaved deciduous forests.Coniferous forests with Picea and Picea-Abies assemblages and variousTsuga species existed at that time. Swamp forests with abundantTaxodium occupied newly formed fresh water lagoons associated withpeat accumulation (Syabryaj and Shchekina, 1983; Syabryaj andStuchlik, 1994; Syabryaj et al., 2007).

The middle Miocene vegetation in the Ukraine Plain wasrepresented by polydominant broadleaved forests with dominantspecies of Quercus, Ulmus, Zelkova, Castanea, Liquidambar, Fagus,Pterocarya, Juglans, and Carpinus. Undergrowth consisted of Corylus,Erica, Lauraceae, Buxus, Ilex, Arecaceae, and Theaceae. Salix and Alnusdominated riparian forest in river valleys and around lakes. Swampforests with Taxodium occurred in the Crimea Peninsula as well as inthe humid continental area. The presence of Keteleeria, Tsuga, Cedrus,Picea, Podocarpus, and Cupressaceae indicates the expansion ofconiferous forests. Additionally, some herb species were also present,but herbaceous assemblages becamemore diverse and dominant afterthe Tshokrakian. A wide river system expanded in the Ukraine Plainduring and at the end of the Karaganian. Due to the late Karaganianregression larger areas of the continental shelf became land, quicklyoccupied by halophilous terrestrial herbs (Chenopodiaceae).

Mixed mesophytic forests with numerous Fagaceae, Juglandaceae,Betulaceae, Aceraceae, and subtropical relicts such as Daphnogene,Alangium, Arecaceae, Sapotaceae, and Myrtaceae were characteristicfor the Volhynian. Coniferous forests covered a large area north of theBlack Sea, while to the east herbaceous communities expanded ondrier soils and replaced the coniferous and mixed mesophytic forests.Open landscapes became a significant feature in the southern andsoutheastern Ukraine. This trend continued in the Bessarabian, whena forest-steppe vegetation existed. The diversity of herbaceous plants

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increased from west to east (Syabryaj and Shchekina, 1983; Syabryajand Stuchlik, 1994; Syabryaj et al., 2007).

4.1.2.2. Bulgaria. A characteristic feature for the Badenian vegetation isthe regular occurrence and abundance of thermophilous taxa likeEngelhardia, Reevesia, Theaceae, Pandanus, Symplocos, Sapotaceae,Araliaceae, Arecaceae, Schizaeaceae, and Gleicheniaceae (Ivanov,1995; Palamarev and Ivanov, 1998; Palamarev et al., 1999; Stuchliket al., 1999; Ivanov et al., 2002; Palamarev and Ivanov, 2004).Temperate elements, such as Fagus, Alnus, Carpinus, Betula, Tilia, andTsuga were less abundant. This suggests the existence of warm,subtropical climate, but a tendency towards slightly cooler conditionsat the end of the period is evident from pollen records (Ivanov et al.,2002, 2007c). Swamp forests also developed. The relief andpalaeogeographic situation controlled the distribution of swampforests (Taxodiaceae, Nyssa, Myrica, Planera) and ecologically relatedriparian forests (Platanus, Planera, Liquidambar, Ulmus, Carya, Pter-ocarya and Salix). The most favourable conditions for these foresttypes existed in the beginning of the late Badenian in NW Bulgaria,probably connected with the so-called “lagoon period” of the DacianBasin (Trashliev, 1984). Macrofloristic data from some middleMiocene fresh water basins in southern Bulgaria reveal some otherpeculiarities of the Bulgarian flora at that time: A high amount ofpalaeotropical elements, such as the high number of Theaceae genera(Eurya, Gordonia, Hartia, Adinandra, Stewartia, and Camellia) in thefossil flora, a significant presence of Lauraceae, Magnoliaceae, andJuglandaceae, dominance of trees and shrubs, a significant proportionof lianas, and, finally, the presence of some relict Paleogene genera(Bozukov 1995; Bozukov and Palamarev, 1995).

At the end of the Middle Miocene (Volhynian and lower part ofBessarabian) typical subtropical elements like Engelhardia, Alangium,Reevesia, Itea, Pandanus, Castanopsis, Sapotaceae, Symplocaceae,Theaceae, Arecaceae tended to decrease in abundance. A similarvegetation change is observed in the Sarmatian of the UkrainianCarpathians (see above) (Syabryaj and Shchekina, 1983). It isaccompanied by an increased significance of arctotertiary elementssuch as Betula, Alnus, Carpinus, Corylus, Fagus, Eucommia, and Tilia inplant communities. The Volhynian vegetation retainsmany features ofthe late Badenian. The distribution of swamp forests in the Volhynianseems to be more restricted when compared to the Badenian.

4.1.2.3. Serbia. The Middle Miocene (Badenian) flora from Serbia iscomprised of numerous laurophyllous elements. Common elementsin mixed mesophytic forests are Laurus, Ocotea, Persea, Daphnogene,Oreodaphne, Laurophyllum, “Ficus”, Magnolia, and certain species ofQuercus (e.g. Q. neriifolia). Typical temperate, deciduous taxa such asSalix, Alnus, Carpinus, Populus, Ulmus, Zelkova, Acer, and some conifersare less diverse and less frequent (Utescher et al., 2007a). Theoccurrence of some legume-like taxa (possibly xerophytes) points tothe presence of dry and open places (azonal vegetation). Riparianforests (Alnus, Populus, Salix, Myrica) were wide spread in rivervalleys.

Floras of the Badenian and Sarmatian are taxonomically verysimilar. However, the most prominent difference between the florasof these two periods is the percentage of deciduous elements, whichgradually increases during the Sarmatian. The Sarmatian palaeobota-nical record is rich and diverse. Important elements in mixedmesophytic forests were laurel species (e.g. Daphogene, Laurus),accompanied by Acer, Betula, Fagus, Carpinus, Zelkova, Salix, Pupulus,Ulmus, Ilex, Prunus, Rhamnus, and Juglans. The occurrence of Ziziphus,Nerium, and legume-type taxa points to a possible seasonality ofprecipitation. Riparian forests (Populus, Platanus, Salix, Myrica,Liquidambar, and Alnus) occupied river valleys and around lakes.Typical swamp forest with dominant Glyptostrobus and Taxodiumwere distributed in flood plains. The presence of Sequoia, Cupressa-ceae, and Pinus suggests existence of mixed conifer-laurel forest under

a warm temperate climate, probably distributed in higher altitudes(Pantić and Dulić, 1992b; Djordjević-Milutinović and Ćulafić, 2008).

4.1.3. The Late Miocene

4.1.3.1. Ukraine. Distribution of swamp forests (Taxodium, Myrica,Osmunda, and Nyssa) decreased in the Ukrainian Carpathians duringthe Pannonian. The riparian forests were widespread at the sametime. Thermophilous plants almost disappeared from the mixedmesophytic forests, and Carya became a dominant plant in thevegetation, together with Juglans, Quercus, Acer, Tilia, and Ulmus.Engelhardia, and Castanea are rare in the pollen spectra. Thefrequencies of Abies and Picea decreased, while different species ofPinus became dominant in coniferous forests. Numerous small freshwater basins with diverse water plant assemblages were typical forthis time.

The vegetation in the Ukrainian Carpathian region changed duringthe Pontian. Picea dominated in coniferous forests in highermountains. The fossil record indicates a lowering of the upper limitof distribution of broadleaved forests in the mountain areas whencompared to the Pannonian. Subtropical species almost disappearedfrom the fossil flora.

In the Ukrainian plain vegetation and climate in the Chersonianwere close to the conditions in the Bessarabian, although somecooling took place, as evident in the palynological data. In thesouthwest, deciduous forests expanded. Woody plant assemblagesopened to the east, and open landscape covered by mesophytic grass-steppe communities appeared. From west to east the role of differentherbs increased, and the composition was enriched. Saline-steppe andsemi-arid desert occupied the southern part of this region. Thereduction of the marine basin and the presence of a wide river systemcaused variable ecological and edaphic conditions, which explainvegetation differentiation.

Different forest communities with Betula, Carpinus Salix, Alnus,Ulmus, Liriodendron, Juglans, Carya, Acer, Aralia, Vitis, Pinus, Picea,diverse herbs, and peat communities were characteristic for theMaeotian. The steppe landscape of the Chersonian was replaced at thevery beginning of the Maeotian by a forest-steppe landscape withopen woody communities. .Small swamp assemblages existed on thewatersheds in the western and central part of the southern Ukraine.Syabryaj et al. (2007) suggest the existence of forest-steppe landscapeand savanna-like plant communities with Acacia, Fabaceae andgrasses. Open and semi-open landscapes with xerophytic plantcommunities predominated till the end of the Maeotian (Syabryajet al., 1993).

A slight increase of humidity in the Pontian influenced thevegetation change for the all of Ukraine. The steppe vegetation hadchanged in character, and Poaceae and other grasses replacedChenopodiaceae. Riparian forests played an important role in rivervalleys. Swamp forests occupied wet areas and coastal regions to thewest, whereas open landscapes with meadows appeared to the east.At the end of the Pontian, forest-steppe landscapes dominated.

4.1.3.2. Bulgaria. The fossil flora from Bulgaria shows a trend todecreasing abundance of palaeotropical elements after the end theBessarabian and throughout the Chersonian. In themesophytic forestsQuercus, Ulmus, Carya, and Pterocaryawere dominant elements, whileSymplocos, Reevesia, Itea, Sapotaceae, and other thermophilousspecies were absent. The swamp forests were reduced in distribution,but the herbs, especially Chenopodiaceae, become more abundant.Other herbaceous plants (Apiaceae, Lamiaceae, Poaceae, Asteroideae,Cichorioideae, Persicaria, Artemisia, and Chenopodiaceae) formedimportant plant communities as well. The maximum distribution ofherbaceous vegetation is recorded in the beginning of the LateMiocene. These higher proportions are combinedwith subxerophytic/xerophytic plants, which became increasingly important — species of

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Celtis, Pistacia, Oleaceae, Ephedra. Some sub-xerophytes were identi-fied in the macrofloral record — Quercus (ex. sect. ilex), Robinia,Arbutus, Berberis, Paliurus, Celastrus, and Caesalpinites (Palamarev,1989, 1990, 1991, Palamarev and Ivanov, 1998, 2004). Thesepalaeobotanical data suggest the appearance of open landscapes andthe development of xerophytic plant communities, which is inaccordance with the data from the Ukrainian Plain. Jacobs et al.(1999) also suggest the steady spread of steppe vegetation and theexistence of a more arid interior in southeastern Europe and Turkey atabout 10 Ma. Fossil phytoliths from Turkey support the assumption ofwidespread herbaceous vegetation during late Miocene (Strömberget al., 2007). The initial open landscapes were of the so-calledparkland type, characterized as mosaic of open areas and smaller orlarger stands of trees. This trend is better expressed in the area of theEuxinian Basin (NE Bulgaria) than in the Forecarpathian Basin (NWBulgaria).

In southern Bulgaria (Sandanski Graben: Ivanov, 2003) diverseherbaceous taxa with relatively high abundances also indicate thepresence of herbaceous communities in open riverine valleys and hillyupland areas. Fossil finds of Hipparion fauna, related to open spacesand steppe or savannah commmunities (Deinoteriums, Mastodonts,Hiparions, Giraffes, Antelopes, Rhynoceroses, Hyaenas e.t.c) alsocorroborate this result (Kojumdgieva et al., 1982; Spassov, 2000;Spassov et al., 2006). The mesophytic forest paleocoenoses weredeveloped on the foothills surrounding the basin, without forming anentire forest belt (see also Palamarev, 1982). The distribution patternof the vegetation was strongly affected by the relief and braided riverswith small ponds and swamps (Spassov et al., 2006).

After this period, during the latest Maeotian and early Pontian,there is again a decline of most herbaceous elements. The swampvegetation spread again along the lakes, and peat bogs were formed inmany areas, e.g. Staniantsi, Beli Breg, Tundzha, Karlovo etc. (Ivanovet al., 2007a,b, 2008; Utescher et al., 2009a). This fact as well as theincrease of riparian vegetation indicates more humid conditions atthat time. The mesophytic forests were more widely distributed.Species of Carya, Fagus, Betula, Quercus, and Ulmus dominated in theseforests, accompanied by Castanea, Carpinus, and Corylus. Somepalaeotropical plants had survived in these communities as relictsfrom middle Miocene vegetation. The dominant vegetation type inhigh mountains was coniferous forests with Pinus, Tsuga, Abies,Keteleeria, Picea, Cedrus, cf. Podocarpus, and Juniperus. Swampvegetation with Alnus, Taxodiaceae (Glyptostrobus, Taxodium), Cyril-laceae, Myrica, Planera, Poaceae, and Cyperaceae, with high propor-tions of ferns (Osmunda and Polypodiaceae) occupied flooded areas inthe proximity of lakes and in swamps. Riparian communities ofPlatanus, Carya, Alnus, Ulmus, Ostrya, Pterocarya, Juglans, Salix,Staphylea, and Liquidambar spread in the river valleys. Communitiesof aquatic plants (Butomus, Potamogeton, Menyanthes, Sparganium,Typha, and Cyperaceae) are also identified. In some cases thevegetation dynamics display hierarchical cyclicity. It is well shownin the section of Staniantsi Basin, where 3 types of cycles are identified(Utescher et al., 2009a). Two of these cycles are expressed byvegetation changes: longer-term cycles (period ~100 kyr) showfrequency oscillations of thermophilous elements and short-termcycles (period ~21.7 kyr), which are expressed as alternation of twotypes of swamp communities (Polypodiaceae-dominated and Osmun-da-dominated). The third cycle is expressed as millennial-scaleclimatic variability (ca. 4.5 kyr, see the section about Late Mioceneclimate below). Such cyclic vegetation and climatic change is notdisclosed from the Serbian and Ukrainian palaeobotanical record, butit is known from northern Greece (Kloosterboer-van Hoeve et al.,2001, 2006) and southwest Romania (Popescu, 2001; Popescu et al.,2006a,b).

In the upper part of the section in the Staniaintsi Basin anexpansion of herbaceous vegetation is observed. The increasingabundance of Asteraceae, combined with a marked decrease in

woody taxa, points to an opening of habitats and most likely a distinctdecrease in mean annual precipitation. This trend is parallel to themolluscan fauna, which yields several terrestrial taxa (Utescher et al.,2009a). An opening of the landscape in the terminal Miocene isrecognized also from palaeofloras from Tundzha and Karlovo Basins inBulgaria and its surroundings (Ivanov and Slavomirova, 2004; Ivanovet al., 2007b).

4.1.3.3. Serbia. During the Pannonian, macrofloras from variouslocalities in Serbia show some drying of the climate when comparedwith the Middle Miocene. This is primarily indicated by a higherrepresentation of Leguminosae and abundant mediterranean taxa.The broadleaved evergreen elements declined, their taxonomicdiversity decreased, and they were mostly dominated by variousspecies of Daphnogene. The deciduous elements were dominated bygenera Ulmus, Zelkova, Juglans, Rhamnus, Carpinus, Cornus, Acer, andFraxinus. The riparian vegetation was dominated by Salix, Alnus, andPopulus, as well as conifers Taxodium and Glyptostrobus. Pinus is moreor less rare except at the Sremska Kamenica locality where it wasamong dominant floral components.

Some of the Pontian sites in Serbia represent diverse, coastalpeatland vegetation with large deposits of brown coal. Swamp forests(Glyptostrobus, Nyssa, Cyrilla, and Myrica), riparian forests (Alnus, Salix,Populus, Platanus,Acer), andherbaceoushygrophytic vegetation (Juncus,Carex, Phragmites, and Typha)werewell developed around the southernmargin of the Pannonian Lake (Pantić, 1989.). Their wide distributionwas forced by the local environment — vast flooded areas and a welldeveloped river system. Palynological data support this assumption andprovide information about wide distribution of swamp and marshvegetation (Pantić andDulić, 1993).Mixedmesophytic forests occupiedmore elevated areas around the water basins and were composed ofdifferent taxa (Magnolia, Laurus, Daphnogene, Carpinus, Ulmus, Juglans,Carya, Pterocarya, Acer, Tilia, Ilex, Sapindus, Bumelia, Vitis, Smilax),mainlydeciduous species, but some evergreen shrubs and trees and lianas alsoparticipated. Thus the vegetation points to a warm and humid climateup to the end of the Pontian. After the Pontian in the Pliocene thevegetation went through significant change. A high number ofthermophilous elements disappeared, while the genera of deciduoustemperate forests became dominant (Fagus, Quercus, Betula, Carpinus,Ostrya, Acer, and Tilia). These changes correspond probably to a slightcooling trend.

4.2. Climate analysis

4.2.1. The Early MioceneClimatic data calculated for the floras discussed above, using the

Coexistence Approach, are shown in Fig. 4. Although the timeresolution that can be obtained from the macrofloral record is limited,major trends and changes of the climate in both Central and EasternParatethys are reflected.

For the early Miocene of the Ukrainian Carpathian region meanannual temperatures (MATs) ranged between 16 °C and 17.5 °C, coldmonth mean temperatures (CMMTs) between 6.5 °C and 8 °C, andwarm month mean temperatures (WMMTs) between 25.5 °C and27.5 °C (Fig. 4). Temperatures tended to be lower in the Ukraine Plain.This was especially true for CMMT, attaining only 2 °C to 7 °C. Climatedata for the Oligocene/Miocene transition in Bulgaria (Babino,Bobovdol Basin) display higher intervals for CMMT (5–12.6 °C),WMMT (25.7–28.1 °C), and MATs 15.6 to 21.1 °C. These values areclose to the temperatures obtained for Ukrainian Carpathians,especially with respect to the lower limits of the coexistence intervals(СА intervals). Palaeoclimate data from Serbia have slightly lowervalues for the temperatures in the latest Chattian (Ravna Reka,Serbia), especially as regards winter temperature— CMMT 2.5–5.8 °C,MATs 13.8–16.6 °C, and WMMT 25.6–25.7 °C. These values are closerto the data obtained for the Ukraine Plain (with CMMT ranging from

Fig. 4. Climatic records for the Miocene of the Central and Eastern Paratethys: [Serbia: (Utescher et al., 2007a), modified; Ukraine (Syabryaj et al., 2007), modified, and Bulgaria: thisstudy)] compared to the continental climatic record of northern Germany (Utescher et al., 2009b) and themarine oxygen isotope record (Zachos et al., 2001); Bo: Bobov Dol / Babino;Ch: Chukourovo; Ru: Ruzhintsi; Ko: Koshava; Dr: Drenovets; Be: Beli Breg.

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2 °C to 7 °C). In the Serbian record, there is a pronounced temperaturedecrease observed. MAT around 15 °C and CMMT around 4 °C areobtained for the Eggenburgian to Ottnangian (e.g. Stamnica flora).

Precipitation records (Fig. 4) indicate that annual precipitationrates (MAPs) are consistently high during the early Miocene in thestudy area, with MAP clearly over 1000 mm: 1122–1206 in theUkrainian Carpathians, 1217–1355 (1146–1146) in the Ukraine Plain,from 1122 to 1355 mm in Babino (Bulgaria), as well as 1003–1355 mm in Chattian and 1122–1194 mm in the Eggenburgian/Ottnangian in Serbia.

4.2.2. The Middle MioceneThe temperature records of Ukraine characterize the Badenian as

warmest period of the whole Miocene, with MAT ranging from 15 °Cto 18 °C and CMMT from 7 °C to 12.5 °C. Again the UkrainianCarpathians appear to be warmer by about 1 to 2 °C when comparedto the Ukraine Plain. The quantitative palaeoclimatic data obtainedfrom palynofloras from Forecarpathian Basin (Bulgaria) displaysimilar climatic conditions for the Badenian (Ivanov et al., 2002).TheMAT coexistence intervals rangemainly between15.6 and 18.4 °C;CMMT intervals between 5.0 and 8.1 °C, and the WMMT coexistence

interval constantly range from 24.7 to 27.8 °C. The highest values forMiddle Miocene annual temperature reach up to17.2–18.4 °C (Ivanovet al., 2002). The data obtained for the Tarkhanian and Karaganian inthe Euxinian Basin (NE Bulgaria) show wider ranges of coexistenceintervals for all climate parameters (MAT 13.6–17.2 °C, CMMT 2.4–7.0 °C, and WMMT 24.7–27.8 °C). This is connected to the lowtaxonomic diversity in these palynofloras (Ivanov et al., 2007c).Data derived from middle Miocene Chukurovo macroflora are in thesame range: MAT 15.9–16.5 °C, CMMT 4.8–5.8 °C, and WMMT 25.8–27.7 °C.

Rising temperatures in the Badenian, as compare to Early Miocene,are detected in the Serbian record. CMMT increased by about 4 °C(Popovac, Slanci, Misaca floras), while the temperature excursionsobserved for MAT and WMMT are only moderate (Utescher et al.,2007a). Although the climatic resolution is delimited in some cases,climatic data are close to those obtained from Bulgarian floras, e.g.MAT 15.6–16.5 °C, CMMT ca. 7 °C, andWMMT 25.6–27.0 °C (Utescheret al., 2007a: Table 3, Slanci). Thus in the whole study area theBadenian constitues a warm climatic phase corresponding to aglobally observed warm time interval, the mid-Miocene ClimateOptimum (Zachos et al., 2001).

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Precipitation records indicate a humid climate for the Badenianperiod in the study area, from Ukrainian Carpathians to the Serbianlakes, with MAP clearly over 1000 mm. MAP coexistence intervalsmost commonly range from 1100 to 1300 mm.

At ca. 14 Ma, a cooling trend is obvious from the temperaturerecords. This cooling, affecting all analyzed variables, is detected in theUkraine Plain and Ukrainian Carpathians. A slight decrease intemperatures is recorded also in Bulgaria in the sediments of theEuxinian and Forecarpathian Basins (Ivanov et al., 2002; Palamarev andIvanov, 2004; Ivanov et al., 2007c) (e.g. core C-12: see Ivanov et al.,2002). Presumably these changes reflect some drying and slight coolingof winter temperatures at the end of the Badenian in Bulgaria, whichprobably corresponds to the beginning of the cooling event about 14–13.5 Ma (Böhme, 2003). It is possible that this event is not welldocumented floristically because of the regression of the ParathetyanSea before the end of the Badenian and lack of fossil record.

Until the end of Middle Miocene (Sarmatian = Volhynian andearly Bessarabian) climatic data in the whole study area do notsignificantly differ from the results obtained for the Badenian. In theUkrainian Carpathians (Sarmatian age) and in the Ukrainian Plain(Volhynian age) climate did not change, and there are no signifi-cant differences. The same is true for the Bulgarian part of theParatethys, where climatic conditions remained relatively stableduring the Volhynian and the lower part of the Bessarabian both inForecarpathian and Euxinian Basins. MAT was 15.6–17.2 °C, CMMTwas mainly 5–7 °C, summer temperatures were at 24.6–27.8 °C andboth CMMT and WMMT show minor oscillations of the upper limit ofcoexistence intervals. The Rouzhintsi macroflora (NW Bulgaria,Volhynian age) displays similar results: MAT 16.5–17.4 °C, CMMTca. 6.2, WMMT 25.9–26.1 °C, with MAP at 1000 mm.

The Serbian palaeoclimatic record shows almost the sametemperatures for the Sarmatian (floras Bela Stena and PancevoBridge). MAT did not exceed 16.6 °C, summer temperatures rangefrom 26.4 to 28.1 °C, and winter temperatures from 5.8 to 7 °C. So, thedata reveal a warm and humid climate up to end of the MiddleMiocene in southeastern Europe, without any dramatic changes (seeIvanov et al., 2002).

4.2.3. The Late MioceneThe late Miocene began at 11.6 Ma (Gradstein et al., 2004b) and

includes the Pannonian and Pontian stages in the Central Paratethysand upper part of the Bessarabian as well as the Chersonian, Maeotian,and Pontian stages in the Eastern Paratethys (Popov et al., 2006; Pilleret al., 2007). It is a period of major vegetational and climatic change inthe study area.

Quantitative data from the Ukrainian floristic record clearlyindicate a climate trend. In the Ukrainian Carpathians a coolingtrend is expressed by all temperature variables analyzed as well asan increase in seasonality of temperature. A considerable decline inCMMT in the Pontian is apparent that could reflect uplift leading toincreasing altitudinal differentiation of the vegetation. In southernUkraine the cooling was accompanied by a gradual drying (Syabryajet al., 2007). The precipitation records indicate that the annualprecipitation rates (MAPs) dropped below 1000 mm in the UkrainePlain. An overall decrease is observed for both the precipitation ofboth the driest and wettest months, more pronounced for the latter(Syabryaj et al., 2007). This aridization trend corresponds to thepartial replacement of forest vegetation by herbaceous communi-ties, which increased in richness and widened from east to west.After the initial cooling during the earlier part of the late Miocene awarming is recorded for the Ukraine Plain at about 7.5 Ma.Especially for upper Maeotian microfloristic record high valuesare obtained, even within the range of Middle Miocene temperatureconditions.

The beginning of the late Miocene also manifests paleoclimaticchanges in the Balkans. Themost extensive climatic record is based on

Bulgarian floras. There a slight decrease in MAT is indicated at the endof the Bessarabian, both in the Euxinian and Forecarpathian basins(Ivanov et al., 2002, 2007c). This period is characterized by lowervalues in almost all climate parameters. MAT coexistence intervalsrange from 13.3 (14.4) to 17.2 °C, corresponding to a decrease of thelower CA limit of about 2 °C as compared to the Bessarabian andVolhynian. A similar cooling is observed in the CA intervals for theCMMT and WMMT. The lower boundary of the coexistence intervalfor summer temperature decreases by about 2 °C, and for wintertemperature by about 3 °C. But themost significant change is reflectedin mean annual precipitation. The coexistence intervals for MAP fallfrom ca. 1076–1308 mm to 652 (740)–759 mm. These climatic datafor the Chersonian indicate that the climate was slightly cooler(particularly in winter) and significantly drier than in the Bessarabian,as is also evident from the vegetation change described above. Theseresults coincide with sedimentological data — chemical precipitationof aragonitic muds at high temperatures and shallow-water condi-tions (Koleva-Rekalova, 1994; Ivanov and Koleva-Rekalova, 1999;Ivanov et al., 2002, 2007c).

Climatic data corroborate the increasingly xerophytic aspect of thevegetation at the end of the Bessarabian and Chersonian in theUkraine plain, as discussed above. These results are well in agreementwith data obtained from small mammals (van Dam, 2006), alsoindicating an aridification trend at the beginning of the late Miocene.Hypsodonty of large mammals based on precipitation maps (Forteliuset al., 2006) suggest that since ca. 11.1 Ma (MN 9 zone) they show ahumidity gradient from continental to more marine conditions fromeast to west. The ecomorphological analyses of large Tortonianmammals (Brayard et al., 2004) and computed ecomorphologicparameters for European large-mammal localities also support theexistence of such an aridity gradient in Late Miocene times, withmoreopen and arid environments in southern Europe.

Quantitative climatic data for the early Maeotian show a slighttrend to a somewhat warmer and more humid climate whencompared to the Chersonian, with MAT=15.6–17.2 °C, CMMT=5.0–6.6 °C, WMMT=24.7–27.3 °C, and MAP up to 1187–1322 mm. Thisearly phase is followed by a cooler period: the lower boundary of MATdecreases by about 2–3 °C, and the lower boundary of CMMT reaches0 °C. WMMT also shows a decreasing trend as expressed by the curvebased on means of coexistence intervals. In the middle part of the laterMaeotian, there is a new shift towardswarmer (MAT, CMMT) andmorehumid conditions, which probably correlates to a warmer episoderecorded for theUkrainePlain at about 7.5 Ma (see above). This period isfollowed by a drier and cooler phase at the endof theMaeotian. Hereweobserve the lowest MAT coexistence intervals, with a lower CA limitbelow 12 °C.

After the cooling in the later Maeotian, the Pontian began with awarming trend. MAT reaches values from 15.6 to17.2 °C, and CMMTfrom 5 to 7 °C. Almost simultaneously MAP increased up to 1187–1308 mm. This event corresponds with the vegetation evolutiondiscussed above, which shows an expansion of swamp and riparianforests during the Lower Pontian. Climatic data reconstructed for BeliBreg Basin (W Bulgaria) also indicate a warm-temperate climate inthe Pontian, with mean annual temperatures around 16 °C, and meantemperature of at least 4 °C during the coldest month. With annualprecipitation rates commonly above 1000 mm, climatic conditionswere overall humid (Ivanov et al., 2007a).

The detailed pollen record (including high-resolution pollenanalysis) from Staniantsi Basin expresses hierarchical cyclicity in thelatest Maeotian/Pontian. Longer-term cycles (period ~100 kyr) showfrequency oscillations of thermophilous elements triggered byclimatic shifts from warmer/wetter to cooler/drier periods. Short-term cycles (period ~21.7 kyr) are expressed as brown-coal/marl/shell layer alternations combined with cyclic change in swampvegetation types. The climatic records reveal a distinct variability intemperature of the warmest month (Utescher et al., 2009a,b). This is

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uncommon in climatic records from central European Cenozoic andpoints to differing sensivities of regional climates.

Although time resolution and number of fossil floras from Serbiaare limited, the climatic data reflect the major trend known as LateMiocene cooling and permit correlations with other continentalclimatic records reconstructed for other realms of the EuropeanCenozoic. For example, CMMT decreased more and earlier in Serbia,pointing to more continental climate conditions during the LateMiocene when compared to Western Europe. But this cooling is ingood accordance with data for the Ukrainian plain and Bulgara. TheSerbian floras indicate the existence of regionally differentiatedprecipitation patterns and specific edaphic conditions. At that time,extended water bodies (Paratethys, Lake Pannon) influenced the localclimate, making it mild and humid. Short-term changes of precipita-tion rates were also possible, similar to changes in the precipitation inBulgaria, but they could not be confirmed because of the imprecisionof available stratigraphical dating. The precipitation rates recon-structed for Serbia tend to be lower than those in the the northwestGerman Cenozoic, but at the same time higher when compared to thelow precipitation rates reconstructed for the beginning of LateMiocene in the area of the Euxinian Basin. Thus this area has anintermediate climate, between the so-called Central European WetZone (Utescher et al., 2007a) and the drier climate of East Ukrainianplane. As obvious from the data calculated seasonally, variableprecipitation rates without summer drought are assumed for localitiesof the neighboring Pannonian area. Also in Serbia a moist type ofclimate existed during the Pontian (Utescher et al., 2007a).

5. Conclusions

The well preserved and diverse palaeofloras from SoutheastEurope (Eastern and Central Paratethys) allow us to reconstructvegetation and climate evolution and to link it with palaeogeographicchanges for the stratigraphic sequence from the Early to LateMiocene.

The Early Miocene flora comprises dominantly evergreen vegeta-tion types. The main vegetation units were subtropical mesophyticbroadleaved forests with, as main components species, of Lauraceae,Juglandaceae, Symplocaceae, Staphyleaceae, Sapotaceae, Theaceae,Arecaceae, Myrtaceae, Gleicheniaceae, Cyatheaceae, Schizaeaceae,Dicksoniaceae etc. Warm-temperate representatives of Juglans, Acer,Tilia, and Quercus were more or less important elements in theseforests. Swamp forests with Glyptostrobus, Taxodium, Comptonia,Myrica, Cyrilla, Andromeda, and Nyssa existed in flooded areas. In theLate Oligocene (Chattian) the presence of some xerophitic taxa pointsto slightly drier conditions (possibly a more pronounced seasonalityof precipitation) in the western part of the study area (Serbia). But atthe end of the Early Miocene broadleaved evergreen generadominated the forests.

In the Early Miocene mean annual temperatures ranged mainlybetween 16 °C and 18 °C, cold month mean temperatures werebetween 6 °C and 8 °C, and warm month mean temperatures —

between 26 °C and 28 °C. The calculations for precipitation showannual total rainfall over 1000 mm. Thus the data indicate a warm andhumid climate.

The middle Miocene fossil record is represented by numerouslocalities in the study area, both with marine/brackish andfreshwater environments. Polydominant multispecies broadleavedforest with many thermophilous taxa like Engelhardia, Reevesia,Theaceae, Symplocos, Sapotaceae, Mastixiaceae, Araliaceae, Areca-ceae, Schizaeaceae, Gleicheniaceae were wide-spread. Significantproportions of Lauraceae, Magnoliaceae, and Juglandaceae arecharacteristic for these plant communities. During the middleMiocene typical subtropical elements tend to decrease in abun-dance. At the same time an increased significance of arctotertiaryelements such as Betula, Alnus, Carpinus, Corylus, Betula, Fagus,Eucommia, Tilia etc. in plant communities is observed. The relief and

palaeogeographic situation controlled distribution of swamp forests(Taxodiaceae, Nyssa, Myrica, Planera), and ecologically relatedriparian forests (Platanus, Planera, Liquidambar, Fraxinus, Pterocarya,Salix). Towards the end of the Middle Miocene open landscapesbecame a characteristic feature in south and southeastern Ukraine.This trend continued in the Bessarabian, when a forest-steppevegetation evolved.

The Middle Miocene was a period of subtropical/warm-temperateand humid climate. The data reveal only small fluctuations in thepalaeoclimatic parameters. The palaeogeographic reorganisation andParatethyan regression led to decreasing MAT and MAP. Thetemperatures tend to decrease over the whole period, probably notmore than 1–2 °C. The climate and vegetation data show a step-by-step transformation without extreme deviations. As a whole theclimate shows a low spatial differentiation, which is in accordancewith conclusions of Bruch et al. (2004) about the absence of wellexpressed latitudinal temperature gradient during the MiddleMiocene.

The beginning of the Late Miocene is marked by significantchanges in the vegetation, forced both by climatic changes andpalaeogeographic reorganizations in the Paratethian realm. Ther-mophilous plants almost disappeared from the mixed mesophyticforests, and Carya, Quercus, Ulmus, Fagus, and Castanea became thedominant arboreal taxa in the vegetation. Palaeobotanical dataindicate the increasingly xerophitic nature of the vegetation at theend of the Bessarabian and Chersonian. Woody plant assemblageshave a more arid character towards the east, and open landscapeappeared. These initial open landscapes were of the so-calledparkland type, a mosaic of open areas with smaller or larger standsof trees. The opening of forest communities was more pronounced inthe east, while in the west herbaceous communities were limited indistribution.

The steppe landscape at the beginning of the Late Miocene wasreplaced at the very beginning of the Maeotian by a forest-steppelandscape with open woody communities in ravines and river valleys,where small swamp assemblages existed on flooded terrains aroundbasins. The vegetation of the later Late Miocene was more differen-tiated in the study area, with less thermophilous plants and a strongdominance of arctotertiary floristic elements.

The beginning of the Late Miocene is marked by a slight coolingand a significant drying of the climate. In the area of the EuxinianBasin the drier period started somewhat earlier, as is indicated alsoby vegetation changes, and the available data suggest drier condi-tions in the Euxinian Basin (Ivanov and Koleva-Rekalova, 1999).These data support the hypothesis about declining precipitation inthe Paratethyan realm (Ivanov and Koleva-Rekalova, 1999; Stuchliket al., 1999; Ivanov et al., 2002). A dry climatic belt was establishednorth or northeast of the Carpathians, gradually spreading to thesouth to the western parts of the Euxinian Basin, later occupying thesouthern parts of the Forecarpathian Basin and the Pannonian Basinto thewest. To thewest dryingwas less intensive. The palaeoclimaticreconstructions of Utescher et al. (2000) for western Germanyimplied that a wet climate existed during the whole Miocene.Probably the climate in the northwest was influenced by theWesterlies and the Atlantic Ocean.

After the initial cooling and drying at the beginning of late Miocene,fluctuations occur in all climateparameters and display cycles of humid/drier and warmer/cooler conditions. These oscillations of MAT, CMMTand WMMT were probably within the limits of 3–4 °C. Together withthe changes in precipitation they were the factors influencingvegetation dynamics. At the end of this period cyclic climatic changeswere recognized too. These changes arewithin the scale ofMilankovitchcycles and probably were orbitally forced.

Thus, the results presented above summarize palaeobotanical dataand draw the general pattern of the vegetational and climaticevolution during the Miocene in southeastern Europe.

273D. Ivanov et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 304 (2011) 262–275

Acknowledgments

We are grateful to H.E. Wright (University of Minnesota, USA) forrevising the English. Our sincere thanks go to anonymous refereeswho greatly improved an earlier version of the manuscript by theircomments and suggestions. This work is a contribution to theinternational program “Neogene Climate Evolution in Eurasia —

NECLIME”. The financial aid of the Projects 436 Bul. 113/139/0-1 / AS103/3-1 (DFG, Germany) and B-1525 (NSF, Bulgaria) is gratefullyacknowledged.

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