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Vegetation and palaeoclimate in the Miocene of Ukraine

S. Syabryaj a, T. Utescher b,⁎, S. Molchanoff a, A.A. Bruch c

a Institute of Geological Sciences NAS of Ukraine, O. Gonchara str. 55-b, Kiev 01601, Ukraineb Geological Institute, University of Bonn, Nussallee 8, D-53115 Bonn, Germany

c Senckenberg Research Institute and Natural History Museum, Senckenberganlage 25, 60325 Frankfurt, Germany

Received 7 March 2005; received in revised form 26 February 2007; accepted 5 March 2007

Abstract

Miocene deposits (marine and terrestrial) distributed throughout the whole territory of Ukraine contain numerouspalaeontological remains including plant macro- and microfossils. There are well known occurrences of fossil leaves as well asfruits and seeds. The Miocene strata of Ukraine belong to the Central and the Eastern Paratethys and deposits from these regionshave been studied palynologically. For the reconstruction of the vegetation of lowland and mountain areas in Ukraine palynologicaldata have been complemented with data obtained from carpological and foliar studies. To obtain quantitative climate datamicrofloral records from the Ukraine Plain and the Ukrainian Carpathians are analyzed using the Coexistence Approach. The maintrends of floral changes were directed by the global climate evolution complicated by regional tectonic processes. Major vegetationchange occurred during the Late Miocene Cooling. In southern Ukraine, the cooling was accompanied by a gradual drying andpartial replacement of forest vegetation by herbaceous communities that increased in richness and expanded from east to west.© 2007 Elsevier B.V. All rights reserved.

Keywords: Palaeobotany; Palynology; Palaeoclimate; Miocene; Ukraine

1. Introduction

Ukraine covers a territory of more than 601 km2 inCentral and Eastern Europe. Its extent from north tosouth is nearly 900 km, and from west to east more than1300 km. The orographic features of Ukraine clearlyreflect the geological structure of the territory. Tectonicmovements, Quaternary glaciations, sea level fluctua-tions of the Black Sea, erosive-accumulative riveractivity, and other physical–geographical processesplayed an important role in forming the contemporaryrelief.

The major part of Ukraine is a wide, uneven plain.Uplands and lowlands are largely underlain by anuneven crystalline basement. The uplands are connectedwith the Ukrainian crystalline shield (from the west toeast: Volynian-Podolian, Pridneprovian, PriazovianUpland and Donetsk Ridge). The main lowlands arePoleskaya, Prichernomorskaya and Pridneprovskaya.The latter, Pridneprovskaya, occupies the middle part ofthe left bank of the Dnieper River. In Ukraine, there aretwo distinct mountain ridges characterized by a steeperrelief and higher altitude when compared to the hillylowlands of the crystalline basement: the Ukrainian orForest Carpathians in the west, and the CrimeaMountains on the Crimea Peninsula. These mountainsbelong to the Alpine orogenic belt.

Palaeogeography, Palaeoclimatology, Palaeoecology 253 (2007) 169–184www.elsevier.com/locate/palaeo

⁎ Corresponding author. Tel.: +49 228 73977; fax: +49 228 739037.E-mail address: utescher@geo.uni-bonn.de (T. Utescher).

0031-0182/$ - see front matter © 2007 Elsevier B.V. All rights reserved.doi:10.1016/j.palaeo.2007.03.038

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The Neogene deposits (marine and terrestrial) areextensive throughout the territory of Ukraine andcontain palaeontological remains including numerousmacro- and microfloras. There are many well-knownoccurrences of leaf remains, most of the Miocene stratahave been studied palynologically. The present work isfocussed on the Miocene vegetation and climate ofmountain and plain areas of Ukraine belonging to theCentral and Eastern Paratethys. Palaeoclimatic recon-structions are based on the palynological record, but allpalynological data have been correlated with the resultsobtained from analysis of macrofloras.

2. Materials and methods

The material for this study is derived from Miocenesediments from two large regions of Ukraine (Fig. 1):the north-eastern part of the Central Paratethys, namelythe outskirts of the Pannonian Basin (UkrainianCarpathian region), and the western part of the EasternParatethys (Ukraine Plain and Crimea). Regional stagesas well as stratigraphic units for each of the aboveregions are shown in Fig. 2.

For palynological analyses the complete stratigraphicsequence of the Miocene deposits is studied: Molassessediments from the Ukrainian Carpathians realm andvarious marine sediments of the Ukraine Plain. In somecases previously published data are used (Shchekina, 1979;Syabryaj and Shchekina, 1983). Each stratigraphic stage ispalaeontologically characterized by foraminifers, ostra-cods (Pishvanova, 1958; Sheremeta, 1958; Venglinsky,1975), and molluscs (Grishkevich and Nevesskaya, 1958).Some stages are dated by large mammal fauna (e.g., Hip-parion fauna; cf. Syabryaj et al., 1993).

All pollen spectra obtained from marine sediments areaveraged because spores and pollen were transported tothe sea or lagoons from upland areas or from the coastalplain, drifted by wind, washed out from soils by rain, andtransported by rivers and streams. Such averaged spectrareflect the composition of plant communities of severalvegetation zones (e.g., all elevations in mountain areas)and from zonal vegetation types originating fromdifferentlatitudes of the Ukraine Plain. The samples analyzed arecollected from cores of numerous wells as well as fromoutcrops (Fig. 1). The numbers of samples evaluated foreach stratigraphic stage is summarized in Table 1.

Fig. 1. Study area with major structural elements and sites analyzed. The site numbers are explained in Table 1.

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To study the palaeoclimate evolution in different partsof Ukraine during the Miocene the CoexistenceApproach (CA) is used. The method is described indetail in Mosbrugger and Utescher (1997). Only a briefexplanation will thus be provided. The distribution ofplant species is – besides other factors – dependent onvarious climatic parameters. The CA follows the nearestliving relative concept. It is based on parameter rangesthat are typical for the extant closest relatives, which areidentified by comparison with the fossil taxa. This isaccomplished by tabulating the climatic conditions of theareas in which these extant representatives exist today.By using this database of extant taxa and their climaticrequirements, “coexistence intervals” of different cli-matic parameters are calculated that allow themajority ofencountered fossil taxa to exist at that location. Theoverlapping of the single taxa ranges defines thecoexistence interval which is then interpreted as therange of the palaeoclimate. The reliability of the resultsobtained increases with the percentage of taxa that may

coexist in this coexistence interval (Mosbrugger andUtescher, 1997). The interval size of the differentparameters decreases in general with the increasingnumbers of species. The CAwas tested extensively withrespect to reliability and resolution, and its advantagesand disadvantages are discussed thoroughly by Mos-brugger and Utescher (1997). For example, it can bedemonstrated that evolutionary shifts of the climateranges of taxa can be detected by statistical methods andby the occurrence of “outliers” which do not fit into thecoexistence interval (Mosbrugger and Utescher, 1997).The core of the Coexistence Approach (CA) isrepresented by the database PALAEOFLORA (Utescherand Mosbrugger, 1990–2007) that includes climate dataof numerous Cenozoic plant taxa based on the climaticrequirements of their nearest living relatives. Initiallyclimate data for all nearest living relatives known for afossil flora are extracted from the database. Thencoexistence intervals are calculating for the selectedclimate variables using an analysis algorithm.

Fig. 2. Regional stages of the Central and Eastern Paratethys according to Harzhauser and Piller (2007-this volume) with stratigraphic units of thePrecarpathian Basin, the Transcarpathian Basin, and the southern Ukraine (after Venglinsky, 1975). Grey: No sedimentary record.

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In the present study, a total of 17 microfloras isanalysed with respect to seven climate variables. Thesevariables are mean annual temperature (MAT), coldmonth mean (CMT), warm month mean (WMT), meanannual precipitation (MAP), precipitation in the warm-est month (MPwarm), precipitation in the driest month(MPdry), and precipitation in the wettest month(MPwet). The microfloras studied each represent asummary for stratigraphical stages and substages, ineach case combined from numerous samples. Toperform the analysis, about 50 additional palynomorphtaxa were entered in the database, as well as climate datasets for about 20 extant relatives potentially providingrelevant climate signals. Single climate variables aresummarized in Tables 2–5. The number of taxa that

contribute to the analysis with climate data ranges from10 to 64 with a mean of 40 taxa. Percentages of extantplant taxa coexisting in the CA climate rangescommonly are well above 90%, frequently even 100%(Tables 2–5). In these cases, the climate data obtainedare considered highly significant (Mosbrugger andUtescher, 1997). In the analyses, no permanent climatic“outliers” are observed. In two cases, data areconsidered less reliable. For the Chaplinka and BolshieKopani floras the calculation of MPwarm providedcoexistence percentages of 88.5% and 80.5%, respec-tively (cf. Table 5). In both cases, however, the numberof taxa pointing to drier conditions equals the number ofthose indicating wetter conditions with respect to theCA results.

Table 1Sites and sample sets analysed in this study

Localities analysed

Region Sites; for site numbers cf. Fig. 1 Sample sets analysed Number of samples

Ukrainian Carpathians Velikaya Began (856 well) (5) Pontian 17 samplesVelikaya Began (856 well) (5) Pannonian 28 samplesBeregovo (632 well) (4) Middle Sarmatian 21 samplesTernova (4u well) (3) Late Badenian 31 samples.Olkhovtsy (outcrops) (3) Early Badenian 30 samplesVoditsa (outcrops) (2) Eggenburgian 27 samplesZaluzh (outcrops) (1) Egerian 28 samples

Ukraine Plain Chaplinka (10k well) (10) Pontian 15 samplesEmetovka (1gk well) (9) Early Maeotian, upper part 26 samplesEmetovka (1gk well) (9) Early Maeotian, lower partEmetovka (1gk well) (9) SarmatianKorobki (40k well) (8) Konkian 38 samplesKorobki (40k well) (8) KaraganianKorobki (40k well) (8) ChokrakianLeniskoe,Crimea (473 well) (7) Tarkhanian 9 samplesBolshie Kopani (2k well) (6) Sakaraulian 19 samplesBolshie Kopani (2k well) (6) Karadzhalganian 18 samples

The locality numbers given are the same as in Fig. 1 showing the geographical position of all the localities. Climate calculations in each case are basedon a sample set. The sample sets are combined according to regional stages (cf. Fig. 2).

Table 2Ukrainian Carpathians, temperatures as calculated by the Coexistence Approach (cf. Syabryaj et al., 2007)

Ukrainian Carpathians, temperature data

Locality name/no. Age Lon. Lat. NT MATn MATx % CMMn CMMx % WMMn WMMx %

VELIKAYA_BEGAN_PO (5) Pontian 22.57 48.23 41 11.6 15.7 97.6 −0.1 1.1 95.1 21.7 24.8 100VELIKAYA_BEGAN_PA (5) Pannonian 22.67 48.23 52 15.2 15.2 96.2 3.8 7 92.3 23 25.5 100BEREGOVO (4) Sarmatian 23 48.25 33 15.7 16.2 100 6.6 7 100 25.4 26.8 100TERNOVA (3) Late Badenian 23.73 48.08 31 15.2 18.4 96.8 6.6 12.5 96.8 25.4 26.8 100OLKHOVTSY (3) Early Badenian 23.7 48.05 32 15.6 18.4 100 6.6 12.5 100 25.4 27.9 100VODITSA (2) Eggenburgian 23.7 48.05 38 16.5 17.6 100 6.6 7.1 100 26 27.5 100ZALUZH (1) Egerian 22.85 48.35 26 15.6 16.1 100 6.6 7.8 100 25.4 25.6 100

NT: number of taxa contributing climate data: MAT: mean annual temperature; CMM: cold month mean; WMM: warm month mean. For eachvariable, minimum (n) and maximum (x) of the coexistence intervals are given. %: Percentages of taxa coexisting; percentages refer to the totalnumber of taxa contributing climate data in the analysis.

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3. Palaeogeography

The general palaeogeographic and palaeobiogeo-graphic development of the Central Paratethys isdescribed by Harzhauser and Piller (2007-this volume).Here, we focus on the special palaeogeographicsituation of the Ukraine territory during the Miocenewhich links the Central and Eastern Paratethys. Thewestern part of Ukraine belongs to the UkrainianCarpathians (Central Paratethys) with the Pre- andTranscarpathian Basins. Basins located in the south,Ukraine Plain and Crimea, belong to the EasternParatethys. For the stratigraphic range of regionalsedimentary units the reader is referred to Fig. 2.

3.1. The Ukrainian Carpathian region

The Inner Carpathian Neogene Basins (Transcarpathianand Precarpathian Basin) are part of the epicontinental areaof the Central Paratethys Sea. During the Miocene thesebasins were connected, and during the early stages ofMiocene, the communication among them was broadenough to let their fauna be very close in systematiccomposition. At the end of the Late Miocene, the seawaysconnecting the basins became narrow and then they closed.Due to fresh water inflow to these basins the fauna changedfrom marine to brackish complexes to freshwater commu-nities (Venglinsky, 1975). The territory of the ParatethysBasin began to reduce at that time, and separate basins, suchas the Pannonian and the Dakian Basin in the west, andEuksinian and Caspian Basin in the southeast, formed. Thelatter were connected by seaways. Temporary connectionsof the Tethys with inflow of open marine water causedchanges of the hydrologic regime that influenced thepalaeocoenosis of the Inner Carpathian Basins.

In the Egerian stage, sediments of the Polyanitskayaand the lower Vorotyshchensksaya suites accumulated

in the shallow Precarpathian Foredeep (Molyavko,1960). The Transcarpathian Basin sediments wererepresented by shallow marine, lagoonal, and continen-tal deposits of the Negrovskaya Suite. Here, normalmarine conditions were not reached until the end of theEgerian (Venglinsky, 1975).

During the Eggenburgian, the Transcarpathian Fore-deep began to form due to increasing subductionprocesses. The ingression of marine water lead to awarm, shallow sea, as indicated by the composition ofinvertebrates (Venglinsky, 1975).

At the end of the Early Miocene, a regression cycletook place and caused profound erosional processes.During the Ottnangian, the re-deposition of continentalsediments formed the Tereshulsky conglomerates andbreccias in the northern part of the TranscarpathianDepression (Venglinsky, 1975).

In addition to the ongoing subsidence, the Karpatianbegan with a global transgression, both of which causedfully marine environments in the Transcarpathian Depres-sion until at the end of the Karpatian (Tereblinsky time), asignificant sea-level drop caused a reduction, isolation, andfinally evaporation of the basin. At the beginning of theBadenian the Alpine orogenesis stimulated the ongoingraising of the Carpathian and Crimea mountains and thesubsidence of the foothills adjacent to the platformmargins. This changed the land–sea distribution in theCentral and Eastern Paratethys areas profoundly incomparison with the Early Miocene. In the Transcar-pathian Depression, volcanic processes dominated thebeginning of the Badenian (Solotvino time; Maleev,1964). Later, during the Teresvenskoe, the most extensivetransgression of the Middle Miocene occurred and formeda basin with normal salinity and a connection to the opensea (Venglinsky, 1975). At the end of Teresvenskoe, theTranscarpathian Basin gradually became isolated. Regres-sion and isolation of the basin finished the Badenian stage.

Table 3Ukrainian Carpathians, precipitation rates calculated by the Coexistence Approach (cf. Syabryaj et al., 2007)

Ukrainian Carpathians, precipitation data

locality name/no. MAPn MAPx % PWARMn PWARMx % PDRYn PDRYx % PWETn PWETx %

VELIKAYA_BEGAN_PO (5) 843 1018 97.6 83 84 95.1 32 41 97.6 109 134 97.6VELIKAYA_BEGAN_PA (5) 1096 1206 94.2 93 107 90.4 32 37 96.2 109 134 94.2BEREGOVO (4) 1033 1281 97 93 131 97 24 41 100 134 139 97TERNOVA (3) 1146 1281 100 94 120 93.5 29 32 96.8 134 153 96.8OLKHOVTSY (3) 1122 1213 100 93 172 100 19 39 100 134 236 100VODITSA (2) 1122 1281 100 118 172 100 26 37 97.4 134 195 97.4ZALUZH (1) 1122 1206 100 99 143 100 32 37 96.2 134 143 96.2

MAP: mean annual precipitation; PWARM: precipitation in the warmest month; PDRY: precipitation in the driest month; PWET: precipitation in thewettest month. For each variable, minimum (n) and maximum (x) of the coexistence intervals are given. %: Percentages of taxa coexisting;percentages refer to the total number of taxa contributing climate data in the analysis.

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In the Precarpathian Depression and on the EasternEuropean platform, the sea broadened its boundariesconsiderably during the Badenian. In the Early Sarma-tian, the Transcarpathian Foredeep was disconnectedfrom the Precarpathian one by the uplifting CarpathianMountains. The beginning of the Sarmatian wascharacterised by intensive orogenic movements, asecond phase of volcanism, and sea level fluctuationsin the inner Carpathian Foredeep (Maleev, 1964). TheEarly Sarmatian regression persisted until the Precar-pathian Foredeep was drained in the Middle Sarmatian.Also the Transcarpathian Basin became shallow;however, a short term transgression of warm waterand the restoration of the connections with theMediterranean Basin took place.

During the Pannonian and the Pontian, the Transcar-patian Foredeep was occupied by fresh water andstrongly influenced by volcanic activity.

3.2. The Ukraine Plain

Tectonicmovements at the boundary of the Paleogeneand Neogene caused uplift of the Ukrainian territory andregression of the Oligocene sea, mainly to the south andthe east (Maleev, 1964). Large parts of the Ukraineterritories adjacent to these uplift areas formed a lowplain where alluvial and diluvial sediments accumulated.The shallow sea in the south covered Crimea, EasternPrichernomor'e, and part of the Ukrainian crystallineshield as a wide gulf and spread into the southern part ofthe Dnieper–Donets Depression. At this time thedeposits of Gornostaevskaya (Karadzhalganian), Cher-nobaevskaya (Sakaraulian), and MayachkovskayaSuites (Kotsakhurian) accumulated.

The regression at the end of Early Miocene displacedthe shoreline further to 48° N latitude. The Tarkhanian,the Nikolaev and Odessa regions, Priazovian Massive,

Table 4Ukraine Plain, temperatures as calculated by the Coexistence Approach (cf. Syabryaj et al., 2007)

Ukraine Plain, temperature data

Locality name/no. Age Lon. Lat. NT MATn MATx % CMMn CMMx % WMMn WMMx %

CHAPLINKA (10) Pontian 33.53 46.37 61 13.8 14.5 91.8 0.9 6.6 96.7 23 24.1 93.4EMETOVKA_E_MAEOT2 (9) Maeotian 30.52 46.75 29 4.9 17.4 96.6 −0.1 10.2 96.6 21.7 27 96.6EMETOVKA_E_MAEOT1 (9) Maeotian 30.52 46.75 10 5.2 13.8 100 −5.9 4.1 100 17.1 24.3 100EMETOVKA_L_SARMAT (9) Sarmatian 30.52 46.75 13 4.6 14.1 100 −7.9 5.8 100 17.7 23.2 100KOROBKI_.UPPER_BAD. (8) Badenian 33.43 45.27 56 15.6 15.6 98.2 6.6 7 94.7 24.7 25.9 98.2KOROBKI._MIDDLE_BAD. (8) Badenian 33.43 45.27 64 15.2 15.6 93.8 4.3 9.7 96.9 25 25.9 96.9KOROBKI_EARLIER_BAD. (8) Badenian 33.43 45.27 55 16.4 16.6 98.2 4.3 10.2 98.2 23.6 25.9 98.2LENINSKOE (7) Tarkhanian 34.03 45.27 54 17 17.1 98.1 6.6 7.1 96.3 24.7 25.9 92.6BOLSHIE_KOPANI_BU. (6) Burdigalian 32.97 46.48 51 15.2 15.6 96.1 2.5 4.1 94.1 23 24.1 94.1BOLSHIE_KOPANI_AQ. (6) Aquitanian 32.97 46.48 41 15.6 15.6 90.2 2.3 7.1 95.1 25.3 26.3 90.2

NT: number of taxa contributing climate data: MAT: mean annual temperature; CMM: cold month mean; WMM (warm month mean). For eachvariable, minimum (n) and maximum (x) of the coexistence intervals are given. %: Percentages of taxa coexisting; percentages refer to the totalnumber of taxa contributing climate data in the analysis.

Table 5Ukraine Plain, precipitation rates calculated by the Coexistence Approach (cf. Syabryaj et al., 2007)

Ukraine Plain, precipitation data

Locality name/no. MAPn MAPx % PWARMn PWARMx % PDRYn PDRYx % PWETn PWETx %

CHAPLINKA (10) 897 1151 98.4 74 80 88.5 41 43 95.1 115 127 96.7EMETOVKA_E_MAEOT2 (9) 735 971 100 47 93 100 9 43 100 90 139 100EMETOVKA_E_MAEOT1 (9) 389 900 100 47 87 100 3 40 100 57 109 100EMETOVKA_L_SARMAT (9) 735 1167 100 72 79 100 29 62 100 90 151 100KOROBKI_.UPPER_BAD. (8) 1304 1356 96.4 82 122 100 42 43 94.6 134 170 94.6KOROBKI._MIDDLE_BAD (8) 1146 1281 96.9 99 111 93.8 26 29 96.9 134 139 95.3KOROBKI_EARL. BAD (8) 1217 1347 96.4 85 111 96.4 24 29 98.2 115 122 96.4LENINSKOE (7) 1146 1322 98.1 82 111 96.3 24 32 98.1 134 139 96.3BOLSHIE_KOPANI_BU. (6) 1217 1355 96.1 85 86 94.1 32 37 98 134 153 96.1BOLSHIE_KOPANI_AQ. (6) 1146 1146 92.7 108 131 80.5 26 29 97.6 108 122 90.2

MAP: mean annual precipitation; PWARM: precipitation in the warmest month; PDRY: precipitation in the driest month; PWET: precipitation in thewettest month. For each variable, minimum (n) and maximum (x) of the coexistence intervals are given. %: Percentages of taxa coexisting;percentages refer to the total number of taxa contributing climate data in the analysis.

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Crimea Mountains, Ukrainian shield, Donbass, theDnieper–Donets Depression, and even the westernpart of Prichernomor'e were above sea level.

Although there is no prominent hiatus observedbetweenTarkhanian and Tshokrakian deposits (Nosovsky,1957), there is evidence from molluscs that in theTshokrakian the rather shallow sea covered wider areasthan in the Tarkhanian (Molyavko, 1960). Further tectonicmovement in the Alpine zone caused a reduction of thebasin at the end of the Tshokrakian. This led to adisintegration of the basin into separate fresh waterlagoons and the erosion of basin sediments.

After a short continental hiatus in Karaganian, thesouthern regions of Ukraine were covered again bymarine water both in the western and eastern parts ofPrichernomor'e. The Karaganian Sea was shallow withlow salinity, probably due to the isolation of the basinfrom the Badenian one of the Central Paratethys. At theend of the Karaganian a minor transgression took place.

The palaeogeographic conditions did not change atthe beginning of the Konkian stage. Only in the secondpart of the Konkian, water of Mediterranean originpenetrated to the Pricdhernomor'e region and changedthe ecological conditions abruptly into normal salinity.At that time, large parts of the Ukraine territory were

covered by two marine basins: the Badenian Sea in thewest and the Konkian Sea in the south which expandedto the Volyn–Podolian region in the west, and to thePriazovian massive in the south.

From the beginning of the Volynian stage, the easternand southern territories of Ukraine were lowered and themarine basin widened. On the western part of Ukrainethe Sarmatian Sea still covered the PrecarpathianForedeep and part of Volyn–Podoliya. It is very possiblethat here the western and eastern basins were connected.In a regressive phase at the end of the Volynian the basinwas closed (Bondarchuk, 1960).

The next transgression from the south took place inthe Bessarabian. Here, the sea penetrated through ancientdepressions onto the terrestrial part (Bondarchuk, 1960).

After the regression at the end of the Bessarabian, thesouthern territory became terrestrial lowland in theKhersonian and the sea remained only in the south-western part of Odessa region, eastern Priazov'e, SteppeCrimea, and the eastern part of Kerchian Peninsula. Inthe Late Khersonian, a new transgression took place,which was less intensive than the Bessarabian one. TheMaeotian transgression from the Tethys was even lessintensive at the beginning of the Pontian the marinebasin in the south of Ukraine reduced once more and its

Fig. 3. Mean annual temperature records for the Neogene of the Ukraine Plane (grey bars) and the Ukrainian Carpathians (black bars). The barsrepresent coexistence intervals as calculated by the Coexistence Approach.

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borders were very similar to those of the extant BlackSea. Although later during the Pontian the marine basinwas widened considerably to the north, the maingeomorphologic elements of the Pontian landscapewere close to Recent ones (Bondarchuk, 1960).

4. Vegetation

Miocene vegetation types and their changes in timeare described here separately for the Ukrainian Car-pathian area and the Ukraine Plain. The vegetationanalysis focuses on interpretation of microfloral recordsobtained from sedimentary successions of the Cenozoicbasins (Fig. 2). Data obtained from macrofloras areconsidered as well, where available.

4.1. The Ukrainian Carpathian region

The polytopic spore and pollen complexes of thePrecarpathian Egerian (Polyanitskaya Suite and lowerpart of Vorotyshchensksaya Suite) and TranscarathianEgerian (Negrovskaya Suite) comprise palynomorphassociations from different altitudinal vegetation zones.At the beginning of the Early Miocene, the Carpathianswere still low, estimated as close to 1000 m in altitude.

With the uplift of the Carpathians, the verticaldifferentiation of plant communities became morepronounced at the end of the Egerian. The compositionof plant communities shows some differences betweenthe inner and outer mountain slopes. This differentiationis observed throughout the Miocene and reinforced inthe Sarmatian. The vegetation represents subtropicalforests and swamp forests with abundant Myricaceaelocally present. Ferns including members of subtropicalfamilies, such as Schizaeaceae, Gleicheniaceae,Cyatheaceae, Matoniaceae and Dicksoniaceae werecharacteristic. On the inner slopes of the Carpathians,mixed coniferous forests and broadleaved forests withsubtropical species of the Juglandaceae family, as wellas chestnut (Castanea) forests were most common(Syabryaj, 1980).

The palynomorph spectra of the Eggenburgianindicate a significant warming during the transgressivecycle of the Eggenburgian. The marine sediments are richin ichtyofauna, foraminifers, ostracods, mollusks, seaurchin needles, otolites and other fish remains, and there isalso evidence for a warm shallow sea (Venglinsky, 1975).In the lowlands, a diverse subtropical forest developed.Here, macrothermal ferns, such as Osmunda regalis, anddifferent species of the Gleicheniaceae and Schizaeaceae

Fig. 4. Cold month and warm month mean temperature records for the Neogene of the Ukraine Plain (grey bars) and the Ukrainian Carpathians (blackbars). The bars represent coexistence intervals as calculated by the Coexistence Approach.

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families coexisted. The plant communities probablyincluded tree-like forms of the Ericaceae and palms inrarer cases. These forests are very similar to extantevergreen forests growing in lowland areas of the Azores.Swamp forests occupied only smaller regions.Apparently,some areas were covered by sclerophyllous Quercuscommunities, which subsequently becamemore importantas a vegetation type. In the broadleaved forests, Engel-hardia, and members of Symplocaceae, Staphyleaceae,Rosaceae and Myrtaceae were present. Temperate taxaoccurred in the mid-elevation and partly in the upperaltitudinal vegetation belts. In the coniferous forests,covering the north-east and south-west slopes, Picea,Abies, Tsuga, and Podocarpus became increasinglyimportant. The latter assemblage can be interpreted as amountain taiga association. In broadleaved forests of theouter slopes, warm temperate genera such as Juglans,Acer, Tilia, and Quercus were abundant. Chestnut forestswere more important when compared to Quercuscommunities. Summarizing up these facts, warm andhumid climate conditions are indicated for the Trans-Carpathian of Ukraine during the Eggenburgian (Syab-ryaj, 1981).

In the regressive cycle of the Ottnangian, no plantremains have been found. In the Late KarpatianVoditskaya Suite we found a poor spore and pollencomplex and there is evidence of an impoverishment in

plant communities with respect to more warm-lovingelements that are found after intensive volcanic activity(Syabryaj, 1991).

In the Carpathian Basin, the Badenian representsanother transgressive cycle (Venglinsky, 1975). Due toan existing seaway between the Central Paratethys andthe Tethys, warm oceanic waters stimulated theexistence of thermophilous terrestrial ferns. Broad-leaved forests expanded, with Ulmus, Castanea, andEngelhardia frequently present. The abundance ofCastanea atavia leaves (Iljinskaya, 1960) recorded inBadenian deposits locally points to the presence ofchestnut forests. As indicated by the co-occurrence ofpollen and leaves Fagus and Liquidambar for the firsttime appeared in the forests of the inner zone of theUkrainian Carpathians. Except for Engelhardia, ther-mophilous species decreased in number. Cinnamomumis reported as a floral element of the lowland forest.

A distinctive flora existed on the outer slopes of theUkrainian Carpathian Mountains where Picea and Ul-mus became more important. In this area, thermophilousspecies disappeared completely pointing to coolerclimatic conditions when compared to the inner zoneof the Ukrainian Carpathians.

In the Late Badenian, thermophilous ferns were stillnumerous. In the Ukrainian Carpathians, humid coastaland riparian forests with Carya, Pterocarya, Liquidambar,

Fig. 5. Cold month mean records reconstructed for Ukraine (cf. Fig. 4) compared to the cold month mean curve of the Neogene Lower Rhine Basin(from Utescher et al., 2000).

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Alangium, and various species of Myrica, less frequentComptonia and Cinnamomum polymorphum (accordingto leaf remains) played an important role (Iljinskaya, 1960;Shvarieva, 1983). The mixed forests included Juglans,Engelhardia, Tilia, Acer, Castanea, Fagus, Quercus, andPlatycarya (the latter is infrequent), as well as Zelkovaand UImus. The shrub layer became more diverse(Lonicera,Diervilla, Elaeagnus,Corylus, and Cornaceae).Subtropical assemblages with Cinnamomum, Myrtaceae,sclerophylous Quercus and thermophilous ferns existed,becoming less frequent towards the end of the Badenian.Increasing percentages of herbaceous pollen, chieflymembers of mountain meadow communities, indicate thepresence of open areas at higher altitudes of the raisingmountains. In the north-eastern part of the UkrainianCarpathians, there is also evidence for subtropical forestcommunities with a similar floral composition, consistent-ly reconstructed from themicro- andmacrofloral record. Inaddition, riparian forests with Alnus were characteristic ofthe outer zone.

The beginning of the Sarmatian was characterized byintensive orogenic movements, a second phase ofvolcanism and a reduced extension of the marine basinsin the Ukrainian Carpathians. The vegetation of theEarly Sarmatian was characterized by coniferous forests

with spruce and spruce-fir assemblages and variousTsuga species. The species T. cf. canadensis may growin coniferous forests and in broadleaved forests as well.The broadleaved deciduous forests were similar to thoseof the Late Badenian. By the end of the Early Sarmatian,pine forests played a major role. Broadleaved forma-tions included beech communities, and, in addition, oak-beech and beech-chestnut assemblages. The resultsobtained from palynological studies accord well withmacrofloral data (Molchanoff, 1999). The broadleaveddeciduous forests were very rich in undergrowth andcontained the last member of Lauraceae family(Cinnamomum) recorded in the Ukrainian Carpathians.

In the middle Sarmatian, shrinking of ocean basinscontinued, fresh water lagoons formed, and peataccumulation began initiating the development ofswamp forests with abundant Taxodium. A short-termtransgression of warm ocean water and restoration of theconnections with the Tethys is indicated by the presenceof Mediterranean marine fauna. The slackening ofvolcanic processes led to warming, which stipulated thereturn of subtropical broadleaved communities withvarious ferns, e.g. Gleicheniaceae, the last palms in theCenozoic of the Ukrainian Carpathians, rare representa-tives of Araucariaceae (Agathis) and expanding areas of

Fig. 6. Mean annual precipitation records for the Neogene of the Ukraine Plain (grey bars) and the Ukrainian Carpathians (black bars). The barsrepresent coexistence intervals as calculated by the Coexistence Approach.

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deciduous forests (Syabryaj, 1991). The Middle Sarma-tian was characterized by a rich mycoflora typical forcoal forming environments.

The Pannonian vegetation changed when comparedto the Middle Sarmatian. Swamp forests with Taxodium,and rarer Osmunda, and Nyssa occupied smaller areasthan in the Middle Sarmatian. In drier stands, Glyptos-trobus occurred. In marginal parts of the swamps,limited Myrica grew. Palmae and Araucariaceae dis-appeared from the shoreline forests and Carya became amain forest forming plant in the vegetation zone of thelow altitudes. In the higher altitudinal levels, severalCarya species, together with Juglans, Quercus, Acer,Tilia, Ulmus, Celtis, rare Engelhardtia, and Castanea,made up an important part of broadleaved forest. Theriparian forests were widespread. In coniferous forests,the frequencies of Abies and Picea decreased whiledifferent species of Pinus became the dominant foresttrees. Numerous small fresh water basins with diversewater plant assemblages were typical for this time.

In the Pontian the vegetation in the UkrainianCarpathian region changed sharply. Coniferous forestsbecame dominant and the role of taiga species in highmountain forests increased. In this period, Picea cf.

excelsa appeared for the first time. The altitudinal limitof broadleaved forests was at lower altitude than in thePannonian. Subtropical species almost had disappeared.

4.2. The Ukraine Plain

The Late Egerian (Karadzhalganian) forest vegetationwas less diverse when compared to the Oligocene as isobvious from the disappearance of ferns belonging to theHymenophyllaceae family, and the impoverishment ofwarm loving and subtropical assemblages (Syabryaj andShchekina, 1983). At that time, representatives ofAraliaceae, Theaceae, Sapotaceae, Symplocaceae, andOleaceae are absent. High percentages of Betula andJuglandaceae in broadleaved forests also indicate coolingat the beginning of the Miocene. Decreasing percentagesof palynomorphs characteristic of laurel forest typevegetation indicate a drying climate.

Due to the transgression of the Maykopian Sea in thesouth of Ukraine in the Sakaraulian the coastalterritories including the lower reach of rivers werecovered by swamp forests with Taxodium, Nyssa, andMyrica frequently occurring in the periphery ofswamp. The river valleys were occupied by broadleaved

Fig. 7. Precipitation records of the driest and the wettest month for the Neogene of the Ukraine Plain (grey bars) and the Ukrainian Carpathians (blackbars). The bars represent coexistence intervals as calculated by the Coexistence Approach.

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forests with abundant Juglandaceae (Juglans, Engel-hardia, and Platycarya) and some representatives of theBetulaceae family (Carpinus, Alnus). Increasing fre-quencies of members of the Palmae, Vitaceae, Theaceae,and Myrtaceae families evidence a warming andmoistening of climate. Acacia appeared for the firsttime. According to Shchekina (1979), this genusapparently grew within open shrubland assemblages,like today in Africa. At the same time, the communitiesof euryphilous and hygrophilous representatives of thetemperate flora became widespread, too. Pollen percen-tages and diversities of conifers decreased whileterrestrial herbs became increasingly important. Accord-ing to Shchekina (1979) all these changes in plant covertestify to an increasingly seasonal distribution ofprecipitation during the year. This stage may becharacterized as a comparatively warm time-interval inthe Miocene.

Due to the regression in the Kotsakhurian time, salinesoils became widespread and were covered by halophi-lous vegetation in which members of Chenopodiaceaepredominated over other herbs, such as Rumex, Poaceae,Asteraceae, and Cyperaceae. The sea regression stimu-lated the lowering of ground water level, which causedthe reduction of Taxodium swamp forest areas. Fresh

water basins were surrounded by meadow vegetationwith representatives of Poaceae, Liliaceae, Ranuncula-ceae, and Caryophyllaceae families and others. Pineforests were not as important as a vegetation type. In thecontinental interior, there were polydominant forests inwhich Quercus, members of Moraceae, and tree-likeand shrub-like plants of the Rosaceae dominated. Tilia,Betula, Fagus, Castanea, Ulmus, numerous trees andshrubs belonging to Fabaceae, Acer, and in single casesMagnolia and Sapindus pollen occur.

With the beginning of the Middle Miocene and theregressive phase of the Tarkhanian, the continental areasof the Ukraine Plain were occupied by pine forests withfour different pine species, Keteleeria, five species ofTsuga, one single species each of Cedrus and Picea,and very rarely Ginkgo. Three species of Podocarpusand Cupressaceae were typical of these forests. In themost humid continental areas and on the CrimeaPeninsula, swamp forests with Taxodium occurred.Polydominant broadleaved forest with dominants ofQuercus, Ulmus, Zelkova, Castanea, Liquidambar, Fa-gus, Pterocarya, Juglans, Carpinus, and Betula occu-pied richer soils. In the undergrowth, Corylus, Erica,Lauraceae, Buxus, Ilex, Palmae, and Theaceae occurred,pointing to the subtropical character of these forests.

Fig. 8. Precipitation record of the warmest month for the Neogene of the Ukraine Plain (grey bars) and the Ukrainian Carpathians (black bars). Thebars represent coexistence intervals as calculated by the Coexistence Approach.

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Salix–Alnus thickets cloaked the river valleys and thelakes. Additionally, many different herb species werepresent.

The findings of large mammal remains (Mastodonangustidens Guv.) in Tshokrakian deposits on the KerchPeninsula are of interest (Borisyak and Belyaeva, 1948).These big animals probably migrated to the Crimea fromthe continent at the time when the Crimea Peninsula wasjoined with the mainland at the end of the first half ofTshokrakian (Molyavko, 1960). The continental vege-tation was similar to that of the Tarkhanian, but lessdiverse. At the end of the Tshokrakian, the role of both,Pinus and Taxodium decreased. Broadleaved forestscovered larger areas, and herbaceous assemblagesbecame more diverse and dominant. At the end of theKaraganian, a wide river system spread in the UkrainePlain. Large watershed areas were occupied by pineforests. Taxodium was less important. Broadleavedforests occupied territories with richer soils. Thesewere mesophyllous, polydominant forests with Quer-cus, Ulmus, Fagus, Zelkova, Alnus, Celtis dominatingand Carpinus, Acer, Carya, Castanea, Fraxinus, Rhus,Nyssa, Liquidambar being less common. The LateKaraganian regression indicated more arid conditions.Climate became more continental. Due to the regressionlarger areas of the continental shelf became land,quickly occupied by terrestrial herbs such as halophilousmembers of the Chenopodiaceae.

During the Konkian – in the territory of contempo-rary Northern Prichernomor'e and further to the north –coniferous forests with different species of Pinusprevailed. Swamp forests with Taxodium nearly dis-appeared. Broadleaved forests were diverse, but someimpoverishment of these communities is observed. Theexpansion of herbaceous communities was charac-terised by the abundance of members of the Chenopo-diaceae family.

In the Volynian, coniferous forests occupied a largeterritory in the south. At the same time, mesophyllousforests with numerous Fagaceae, Juglandaceae, Betula-ceae, Aceraceae, and subtropical relicts such as Cinna-momum, lianas (e.g., Alangium) were characteristic. Tothe east, herbaceous communities and meadows spreadon drier soils and replaced the coniferous and mixedforests. Open landscapes became a significant feature inthe southern and south-eastern Ukraine.

In the Bessarabian, a forest steppe vegetation with atypical fauna existed: Mastodons, giraffes, poll-rhinoc-eroses (Dnepropetrovsk region) and hipparions (Zapor-ozh'e region). The remains of beaver and plant remainstestify the presence of a river net and wide river valleys.In the southern region, the Early Bessarabian flora and

vegetation shared the features of the Early Sarmatian.Various broadleaved forest associations with subtropicalelements (Palmae, Sapotaceae, Myrtaceae) existed. Thespatial differentiation, especially in the southern part,continued. From west to east, the diversities of differentherbaceous species increased. Halophytic assemblagesoccupied the whole Azov coast, and meadow commu-nities existed in wetter places.

In the Khersonian continental deposits, a rich Hip-parion fauna (hipparions, gazelles, poll-rhinoceros,giraffes, mastodons) is preserved. Vegetation andclimate were similar to those of the Bessarabian,although some cooling took place almost over thewhole Ukraine. In the Southwestern Ukraine, deciduousforests spread. Towards the east, woody plant assem-blages became more open; the forests were interspersedand combined with open landscape towards the south.The latter were covered by a mesophytic gramineousmotley grass-steppe community, a diverse association ofspecific herbs and grasses. The salty steppe and semi-arid deserts occupied the southern part of this region.The existence of various different ecological andedaphic conditions, the evolution of the marine basinand the presence of a wide river system may explainthese differentiations.

According to macrofloral data (Pimenova, 1937;Dorofeev, 1955) a turgai type flora was characteristic forthe Maeotian. According to palynological studies,different forest communities with Betula, CarpinusSalix, Alnus, Ulmus, Liriodendron, Juglans, Carya,Acer, Aralia, Vitis, Pinus, Picea, diverse herbs, andpeat communities were characteristic. The steppelandscape of the Khersonian was replaced at the verybeginning of the Maeotian by a forest-steppe landscapewith small woody communities in ravines and rivervalleys, where small swamp assemblages existed on thewatersheds in the western and central part of theSouthern Ukraine. The forest-steppe landscape withAcacia and other genera of the Fabaceae and Poaceaefamilies, as well as motley grass prove the existence ofsavanna-like communities. Halophytic communitieswith abundant Chenopodiaceae became widespread asa result of marine regression. At the end of the Maeotian,open and semi-open landscapes with more xerophyticplant communities predominated, and the number ofanimal inhabitants of forest biotopes considerablydecreased (Syabryaj et al., 1993).

In the Pontian, continental deposits fresh waterfishes, fresh water molluscs, tortoises, birds and landmammals (Hipparion fauna) occurred. The land border-ing the Pontian Sea was a low-lying plain. Thegeomorphology was similar to the present. The steppe

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vegetation had changed in character. Poaceae andmotley grass replaced the numerous Chenopodiaceaeand valley forests played an important role. At the end ofthe Pontian, forest-steppe landscapes dominated. Taxo-dium swamp forests occupied wet areas of the westernpart of the Prichernomor'e whereas in the eastern part,plant communities of open landscapes with meadowsappeared. A slight increase of humidity in the Pontianwas characteristic for the whole Ukraine. In coastalregions of the western Prichernomor'e, swamp forestsdevelopped.

5. Palaeoclimate

For the data calculated by the Coexistence Approach(Tables 2–5) climate records can be plotted for theUkrainian Carpathian region and for the Ukraine Plain(Figs. 3–8). Although time resolution of these records islimited, trends and major changes of the climateevolution in both regions are reflected and can bequantified.

As is obvious from the temperature ranges on Figs. 3and 4, temperatures were significantly higher in theUkrainian Carpathians than in the Ukraine Plain duringthe Early and Middle Miocene. In the Late Miocene, incontrast, temperatures were about at the same level inboth regions or even tended to be higher in the UkrainePlain. For the Early Miocene of the UkrainianCarpathian region mean annual temperatures (MATs)ranging between 16 °C and 17.5 °C, cold month meantemperatures (CMTs) between 6.5 °C and 8 °C, andwarm month mean temperatures (WMTs) between25.5 °C and 27.5 °C are obtained. WMT calculated forthe Eggenburgian is among the highest observed in theUkrainian Carpathian record. In the Early Miocene,temperatures tended to be lower in the Ukraine Plain.This was especially true for CMT attaining only 2 °C to7 °C. In temperature records of both regions, highestvalues result for the Badenian, with MAT rangingbetween 15 °C to 18 °C and CMT between 7 °C and12.5 °C. Again, the Ukrainian Carpathians appear to bewarmer by about 1 °C to 2 °C when compared to theUkraine Plain. At ca.14 Ma, a cooling trend is obviousfrom the temperature records. This cooling, affecting allvariables analysed, can be detected in both geo-tectonicregions — Ukraine Plain and Ukrainian Carpathians. Inthe Ukainian Carpathians, temperatures show a declin-ing trend. In the Pontian, there is a considerable declinein CMT observed, nearly attaining freezing point, whileWMT declined more steadily (Fig. 4). Thus, the LateMiocene Cooling was connected to a considerableincrease in seasonality in this region. Different condi-

tions are observed for the Ukraine Plain There, awarming is recorded at about 7.5 Ma, subsequent to theinitial cooling during the earlier part of the LateMiocene. Especially for the later Maeotian microfloralrecord of Emetovka (Table 1) high values are obtained,being even within the range of Middle Miocenetemperatures. In the Ukrainian Carpathians, this warm-ing trend in the upper part of the Late Miocene is absentor cannot be resolved by the data, while in the UkrainianCarpathians, seasonality in temperature increased to-wards the top of the Late Miocene although the meanannual range of temperature did not undergo muchchange in the Ukraine Plain at that time. However,temperature records point to greater climate variabilityduring the Late Miocene when compared to theCarpathian realm (Fig. 4).

Precipitation records (Figs. 6–8) indicate that annualprecipitation rates (MAPs) are consistently high duringthe Early and Middle Miocene in the UkrainianCarpathians, and in the Ukraine Plain as well, withMAP clearly above 1000 mm. However, from theSarmatian and the Pannonian palynological record ofthe Ukraine Plain, significantly lower rates (b1000 mm)are evident. As shown by the record of the UkrainePlain, precipitation rates in the driest (MPdry) and thewettest month (MPwet), respectively, peak in theBadenian and the Sarmatian. During the Late Miocene,there is an overall decrease observed for both, driest andwettest month. This decrease is more pronounced in thewettest month precipitation than in the driest monthprecipitation, but in the Late Miocene (Maeotian) of theUkrainian Plain, MPdry rates of almost 0 mm arepossible, according to the resulting coexistence inter-vals. In the Ukrainian Carpathians, MPdry undergoes nochanges during the Late Miocene and stays consistentlyat 40 mm. However, there is a slight decrease observedfor MPwet. Precipitation rates of the warmest month(Fig. 8) show an overall decreasing trend in the time-span from Early to Late Miocene. This trend is wellreflected in both records of Ukraine. For the UkrainePlain, however, drier conditions are indicated for thelater Early Miocene and the Pannonian, when comparedto the Ukrainian Carpathians. The rates of summerrainfall tend to be lowest in the Pannonian of theUkraine Plane, but still were above 50 mm.

6. Discussion and conclusions

The vegetational evolution in the Ukrainian Car-pathians and the Ukraine Plane during the Miocene notonly reflects global climate change but also stronglydepends on palaeogeographic settings, orogenic

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processes and uplift, and significant movements oftectonic plates. As shown by the analyses of palyno-morphs and the macrofloral record, forest vegetationdominated the lowland and upland areas of theUkrainian Carpathians during the Miocene, formingdifferent altitudinal vegetation zones. As a result ofstronger uplift, an open alpine belt existed in theBadenian. In the Southern Ukraine, mixed forests rich inmesophyllous elements predominated during the Earlyand Middle Miocene. The role of xerophilous elementsincreased over time, and, from the Sarmatian onward,the landscape became more open. As is obvious fromthe palynomorph records, a progressive drying andcooling characterized the Late Miocene causing amigration of steppe landscapes from the east to the west.

In the Ukrainian Carpathians, the overall highertemperatures during the Early and Middle Miocene,when compared to the Ukrainian Plain, coincide wellwith the warmer aspect of the vegetation typesreconstructed for this area (chapter 3.1). One causemay be the more southerly latitudinal plate position ofthe Ukrainian Carpathian region during most of theMiocene. The strong decreasing trend of cold monthmean temperature indicated for the Late Miocene mostprobably is connected to a significant northward shift ofthe tectonic plate. The major vegetation changesobserved during the Pontian of the Ukrainian Car-pathians, like the dominance of coniferous forest and thedisappearance of subtropical elements, are correlatedwith a decrease in winter temperature from 5 °C in thePannonian to almost 0 °C. As described earlier, plantassociations reconstructed for the Ukrainian Carpathiansgenerally have a humid character. Correspondingly,mean annual precipitation calculated by the CA is wellabove 1000 mm except in the Pontian, and precipitationrates of the driest month range between 25 to 45 mmthroughout the Miocene.

In the Ukrainian Plain, the time-span between theEgerian and the Tarkhanian is characterized by thepresence of thermophilous forest associations. Thiscoincides with the reconstructed temperatures withmean annual temperature not below 15 °C, and coldmonth mean ranging between 2 °C and 10 °C whiletemperatures calculated for the Tarkhanian stage areamong the highest. Minor temperature fluctuations, aspartly indicated by changing vegetation, are not resolvedin the climate record. All the associations observed in theabove time-span have a dominantly humid character; insome cases, a more pronounced seasonality of precip-itation can be assumed. The moistening trend in the earlypart of the Miocene, from the Karadzhalanian to theSakaraulian, as derived from the floral record, is also

obvious from the precipitation curves (especiallyprecipitation of the wettest month, cf. Fig. 7). As evidentfrom the interpretation of palynological assemblages, theonset of cooling in the Tschokrakian/Karaganian (cf.cold month mean record in Fig. 4) is connected to asubsequent drying and increasing continentality ofclimate (cf. chapter 3.2). In the precipitation curves,rates have clearly decreased with the beginning of theBessarabian. According to the CA results, this evolutionculminates in the earlier Maeotian where driest monthprecipitation rates near 0 mm and a mean annualprecipitation rate of 400 mm at a minimum are notcontradicted by the data. However, the warmest monthobviously was not identical to the driest month (cf.Figs. 7; 8). Data from vegetation analyses and CA resultsconsistently indicate higher precipitation rates againduring the Pontian.

When comparing the climate records reconstructedfor the Ukrainian Miocene (Figs. 3–8) with globaltemperature evolution, both areas, the UkrainianCarpathians and the Ukraine Plain, show that theBadenian is identified as the warmest interval. Thiswarm time-interval corresponds to the “Mid MioceneClimate Optimum” (Zachos et al., 2001). In bothtemperature records, the cooling trend connected tothe expansion of polar ice-sheets (e.g., Zachos et al.,2001) starts at ca.14 Ma (Figs. 3; 4) as is described fromother continental areas of Western Eurasia (e.g., LowerRhine Basin, W Germany; Utescher et al., 2000). Highertemperatures observed in the Pontian of the UkrainePlain climate record (Figs. 3; 4) are also observed inroughly time-equivalent horizons of the Lower RhineBasin. There, the temperature excursion has beencorrelated to the warming succeeding the MTi-4 isotopeevent (cf. Abreu et al., 1998).

As indicated for the Central European Cenozoicclimate evolution (Utescher et al., 2000; Mosbrugger etal., 2005), the cooling starting at the end of the MiddleMiocene is dominantly controlled by decreasing wintertemperatures and increased seasonality of climate.When comparing Miocene temperatures for NorthwestGermany (Lower Rhine Basin; Utescher et al., 2000)and Ukraine, mean annual temperatures calculated forcorresponding time-intervals are roughly equivalentwhile in the obviously oceanic-influenced LowerRhine Basin cold month means are higher by an averageof 2.5 °C (Fig. 5). Comparable CMTs, however, arereported from time-equivalent records from the CentralParatethys (Mosbrugger et al., 2005).

In the present study, the main trend of the floral andvegetation change in Ukraine was correlated with theglobal climate evolution. Major vegetation changes are

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directed by global climatic development, e.g., the MidMiocene Climatic Optimum and the Late MioceneCooling. This process was complicated by tectonicdevelopment. The tectonic influence was stronger inmountain regions. On this background of global trends,smaller-scale climate oscillations of different durationand intensity also took place.

Acknowledgements

This work is a contribution to the program “NeogeneClimate Evolution in Eurasia — NECLIME”. Theauthors thank all colleagues and NECLIME membersfor fertile discussions. We like to thank E. Zastawniakand R. Burnham for carefully reviewing our manuscriptand for their helpful comments. The financial aid of theGerman Science Foundation (DFG) is gratefullyacknowledged.

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