Palaeoclimate and vegetation change in Serbia during the last 30 Ma

This article was published in an Elsevier journal. The attached copyis furnished to the author for non-commercial research and

education use, including for instruction at the author’s institution,sharing with colleagues and providing to institution administration.

Other uses, including reproduction and distribution, or selling orlicensing copies, or posting to personal, institutional or third party

websites are prohibited.

In most cases authors are permitted to post their version of thearticle (e.g. in Word or Tex form) to their personal website orinstitutional repository. Authors requiring further information

regarding Elsevier’s archiving and manuscript policies areencouraged to visit:

http://www.elsevier.com/copyright

Author's personal copy

Palaeoclimate and vegetation change in Serbia during the last 30 Ma

T. Utescher a,⁎, D. Djordjevic-Milutinovic b, A. Bruch c, V. Mosbrugger c

a Geological Institute, University of Bonn, Nussallee 8, 53115 Bonn, Germanyb Natural History Museum, Njegoseva 51, 11000 Belgrade, Serbia and Montenegro

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

Received 9 February 2005; received in revised form 26 February 2007; accepted 5 March 2007

Abstract

In the present study, 14 published megafloras from the Serbian Cenozoic are analyzed with respect to vegetation type,palaeoclimate, and palaeogeographic settings. The floras cover a time-span from the Oligocene to the late Miocene. The resultsobtained are compared with continental climate records from other parts of Europe and discussed in the context of global climatechange. To obtain a quantitative palaeoclimate record a total of seven different climate variables is calculated for each of the florasusing the Coexistence Approach. It is shown that basic patterns of vegetation change, such as the immigration of arcto-tertiary,deciduous floristic elements during the early Miocene and the decreasing diversity of laurophyllous taxa during the late Miocene,are ruled by climate change and a changing palaeogeography. The Serbian Cenozoic climate record shows a steep cooling gradientduring the Oligocene that is most probably is connected to a northward movement of tectonic plates. The globally observed MiddleMiocene Climate Optimum and the Late Miocene Cooling are well reflected. According to the palaeotemperatures calculated awarm temperate climate existed in Serbia throughout the time-span examined. The Late Miocene Cooling is most pronounced inwinter temperatures and is connected to increasing seasonality. Precipitation rates obtained for the Serbian Cenozoic, especiallythose of the warmest and wettest months, tend to be lower when compared to the Central European Cenozoic. According to climateanalysis and the interpretation of vegetational data there is evidence for regionally drier conditions and increased seasonality ofprecipitation in the time-span from the late Badenian to the early Sarmatian.© 2007 Elsevier B.V. All rights reserved.

Keywords: Cenozoic; Serbia; Palaeobotany; Megafloras; Palaeoclimate; Fixed floras

1. Introduction

The present analysis is based on 14megafloras from theSerbian Cenozoic. 12 of these floras have been studied byNikola Pantic. Pantic spent many years working on theCenozoic floral record in several regions of formerYugoslavia, especially in Serbia. His interest also focusedon problems concerning palaeotectonics and biostratigra-

phy. His extensive research resulted in establishing the so-called Fixed Floras of Serbia, characteristic floras withprecisely determined ages that can be studied by a variety ofcomparativemethods (e.g., Pantic, 1956, 1984;Mihajlovic,1977; Pantic and Mihajlovic. 1977; 1985). In the presentstudy, these floras and additional data from other localitiesmore recently published are analyzed with respect topalaeoclimate and vegetation change in the SerbianCenozoic during the last 30 Ma, under the constraints ofa changing palaeogeography, in order to improve ourknowledge about a key area at the transition between theTethys and Paratethys.

Palaeogeography, Palaeoclimatology, Palaeoecology 253 (2007) 157–168www.elsevier.com/locate/palaeo

⁎ Corresponding author. Tel.: +49 228 739773; fax: +49 228 739037.E-mail address: [email protected] (T. Utescher).

0031-0182/$ - see front matter © 2007 Elsevier B.V. All rights reserved.doi:10.1016/j.palaeo.2007.03.037

Author's personal copy

Serbia, as a part of the Balkan Peninsula, represents anewly formed continental area connected to the southernpart of Palaeo-Europe since the late Oligocene (Pantic,1984). In the Cretaceous and the Palaeogene, the present-day Balkans represented an archipelago within the TethyanOcean. During the late Oligocene, the island arc collidedwith Palaeo-Europe and the Paratethys formed. TheParatethys existed during the whole of the Neogene, eitheras a marine environment characterized by a complexpattern of changing seaways and landbridges or as afreshwater lake (e.g., Rögl, 1998, 1999; Popov et al., 2004).The palaeogeographic evolution of the study area isoutlined in Harzhauser and Piller (2007-this volume). In

the Neogene, the presence of the vast water bodies in theParatethys realm had a huge impact on palaeoclimate andvegetation in Serbia.

Due to the specific palaeogeographic situation, thegeomorphological evolution, and the comparativelysouthern latitudinal position of the Balkans, the floristicand physiognomic composition of the palaeofloras ofSerbia significantly differs from contemporaneouspalaeofloras of Central and Western Europe. Especiallyfrom the middle Miocene on, “Mediterranean” and“subtropical”, warmth-loving floristic elements, such asLaurophyllum type, Daphnogene, and evergreen Quer-cus species reach a higher importance in the floral spectra

Fig. 1. Geological sketch map showing the outline of Cenozoic deposits in Serbia with the locations of the floras indicated. For details of floras seeTable 1.

158 T. Utescher et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 253 (2007) 157–168

Author's personal copy

when compared to Central Europe. Consequently, thispart of the Paratethys has been described as a refuge areafor the dominantly evergreen vegetation, e.g. the so-calledYounger Mastixoid Floras, widely distributed all overCentral Europe in the early to middle Miocene (e.g., Mai,1995; Kovar-Eder et al., 1996; Kovar-Eder, 2003). This isalso true for the present vegetation of the Balkans, namelyfor its south-eastern part, where late Cenozoic andPleistocene relicts may reach considerably higher percen-tages when compared to the vegetation of Central Europe.However, these special conditions prevailing in Serbiaduring the Cenozoic are not really quantified bypalaeoclimate data so far. The occurrence of sclerophyl-lous and legume-like taxa in the Miocene floral record ofSerbia and their climatic implications have been widelydiscussed and were referred to drier climate conditions(e.g., Palamarev, 1989; Mai, 1995; Kovar-Eder, 2003).However, the degree of drying is still a matter of debatebecause quantitative precipitation data were not available.To improve our knowledge, quantitative climate data arederived from the palaeobotanical record of Serbia and arediscussed in the context of vegetation change andchanging palaeogeographical boundary conditions.

2. Materials and methods

All the Cenozoic floras analyzed in the present studyoriginate from localities south of the Danube nearBelgrade, within a maximum distance of about 150 km

from the capital (Fig. 1). The 14 floras are compiledfrom literature and listed in Table 1, with referencescited. Details on stratigraphical ages and method ofdating are given in Table 2. The Oligocene to lateMiocene floras cover a time-span of almost 30 Ma.Important criteria for selecting the floras were theavailability of stratigraphical data, as well as a high

Table 1Serbian megafloras analyzed in the present study

Serbian floras analyzed

Flora Ntaxa Reference

Bogovina 14 Pantic, 1956;Pantic, 1984

Ravna Reka 45 Pantic, 1956Stamnica 31 Pantic, 1956Popovac 23 Pantic, 1956Slanci 35 Mihajlovic, 1978Misaca 22 Pantic, 1956Seliste 36 Milovanovic and

Mihajlovic, 1984Bukovac (Saranovo) 30 Pantic, 1956Bozdarevac 37 Stevanovic and Pantic, 1954;

Pantic, 1956Bela Stena 33 Pantic, 1956Pancevo Bridge 41 Pantic and Mihajlovic, 1977Dubona II 42 Pantic, 1956; Pantic and

Mihajlovic, 1977Crveni Breg Grocka 31 Pantic, 1956, Mihajlovic and

Lazarevic, 1999Osojna (Kladovo) 22 Pantic, 1956

Ntaxa: number of taxa.

Table 2Serbian megafloras analyzed in the present study

Stratigraphical data

Flora Stratigraphicalage

Dating method;floral complexafter Mai (1995)

Reference

Bogovina Rupelian (?);Chattian (?)(MP24)

Mammaldating;Kiscellcomplex

Mai, 1995;Pantic, 1956;Pantic, 1984

Ravna Reka Latest Chattian(MP30)

Mammaldating

Mai, 1995

Stamnica Eggenburgian,Ottnanigian

Pantic, 1956

Popovac Badenian(earlier part)

Wieliczka–Viehausencomplex

Pantic, 1956;Mai, 1995

Slanci Badenian (?) Mihajlovic,1978

Misaca Badenian(later part)

Wieliczka–Viehausencomplex

Pantic, 1956;Mai, 1995

Seliste Badenian toEarly Sarmatian

Freshwatermolluscs

Milovanovicand Mihajlovic,1984

Bukovac(Saranovo)

Late Badenianto EarlierSarmatian

Stare Gliwice–Unterwohlbachcomplex

Pantic, 1956;Mai, 1995

Bozdarevac Late Badenianto EarlierSarmatian

Stare Gliwice–Unterwohlbachcomplex

Stevanovic andPantic 1954;Pantic, 1956;Mai, 1995

Bela Stena EarlySarmatian,later part

Erdöbényecomplex

Pantic, 1956;Mai 1995

PancevoBridge

Sarmatian Marine fauna Pantic andMihajlovic,1977

Dubona II LaterPannonian

Vösendorf–Rószaszentmártoncomplex

Pantic, 1956;Pantic andMihajlovic,1977; Mai, 1995

Crveni BregGrocka

LaterPannonianto Pontian

Vösendorf–Rószaszentmártoncomplex

Pantic, 1956;Mai, 1995;Mihajlovic andLazarevic, 1999

Osojna(Kladovo)

LaterPannonianto Pontian

Vösendorf–Rószaszentmártoncomplex

Pantic, 1956;Mai, 1995

Stratigraphical ages and dating method.

159T. Utescher et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 253 (2007) 157–168

Author's personal copy

diversity of the floral record. The floras comprise 14 to42 different fossil taxa, at a mean diversity of about 30taxa (Table 1). For each of the localities analyzed,composition of the flora, vegetation type, and palaeo-geographic constraints are briefly outlined.

To obtain quantitative palaeoclimate data that can becompared to the results obtained from other continentalregions (e.g., Mosbrugger et al., 2005) the CoexistenceApproach (CA) is used (Mosbrugger andUtescher, 1997).The CA follows the nearest living relative concept. It is

based on parameter ranges that are typical for theconsidered plants that are identified as representatives ofthe fossil taxa. This is accomplished by listing the climaticconditions of the areas in which these extant representa-tives exist today. By using a database of extant taxa andtheir climatic requirements, ‘‘coexistence intervals’’ ofdifferent climatic parameters can be calculated thatallowed the majority of considered plant taxa to exist atthat location. The overlapping of the single taxa rangesdefines the coexistence interval which represents the

Table 3Temperature data calculated by the CA (cf. Utescher et al., 2006)

Temperature data

Flora Nt MATn MATx % CMMn CMMx % WMMn WMMx %

Bogovina 9 17.9 21.3 88.9 7 13.3 88.9 26.4 28.1 88.9Ravna Reka 29 13.8 16.6 100 2.5 5.8 96.6 25.6 25.7 96.6Stamnica 21 14.4 16.6 95.2 3.7 3.8 95.2 25.6 27 95.2Popovac 15 14.4 21.3 93.3 3.7 13.3 93.3 27.2 28.1 100Slanci 25 15.6 16.5 92.0 7 7 96 25.6 27 96Misaca 14 14.4 16.6 100 5.6 11.7 100 25.7 28,1 100Seliste 27 13.5 16.5 100 3.8 7 100 25.7 27 100Bukovac (Saranovo) 23 14.4 15.4 95.7 3.7 5.8 95.7 25.7 26.4 95.7Bozdarevac 27 13.8 17.3 92.6 5.6 7 88.9 26.5 26.7 96.3Bela Stena 18 15.6 16.6 83.3 5 7 88.9 27.9 28.1 94.4Pancevo Bridge 15 14.1 16.6 100 5.8 7 100 26.4 28.1 93.3Dubona II 20 14.4 15.4 100 3.7 4.8 100 25.7 26.4 100Crveni Breg Grocka 28 13.8 15.4 96.4 −0.1 4.1 100 25.7 26.4 96.4Osojna (Kladovo) 19 13.8 16.6 94.7 1.8 5.8 94.7 26.5 26.7 100

Nt: number of taxa contributing climate data; for each parameter the minimum and maximum of the coexistence interval and the percentage of coexistingtaxa are given.

Table 4Precipitation data calculated by the CA (cf. Utescher et al., 2006)

Precipitation data

Flora MAPn MAPx % PWARMn PWARMx % PDRYn PDRYx % PWETn PWETx %

Bogovina 867 1384 100 81 96 100 32 46 100 160 163 100Ravna Reka 897 975 96.6 84 86 100 43 43 100 135 143 96.6

1003 1355Stamnica 1122 1194 100 90 96 95.2 42 59 95.2 131 159 100Popovac 1255 1613 100 90 196 100 42 43 93.3 131 180 93.3Slanci 823 1237 100 79 93 96 43 61 96 131 139 96Misaca 867 1018 93.3 90 177 86.7 42 62 93.3 131 164 100

1122 1187Seliste 1194 1355 96.3 105 113 92.6 32 43 96.3 116 170 92.6Bukovac (Saranovo) 1122 1237 100 58 94 100 42 43 100 131 134 100Bozdarevac 867 1018 96.3 89 94 92.6 11 20 92.6 125 151 100

32 63Bela Stena 1255 1356 100 90 113 94.4 42 43 100 131 151 88.9Pancevo Bridge 867 1032 100 81 92 100 41 59 100 125 130 100Dubona II 1122 1237 100 90 94 90 42 47 100 131 134 100Crveni Breg Grocka 897 1297 100 84 84 89.3 32 47 92.9 131 170 96.4Osojna (Kladovo) 897 1297 100 84 84 100 42 59 100 125 151 100

Nt: number of taxa contributing climate data; for each parameter the minimum and maximum of the coexistence interval and the percentage of coexistingtaxa are given. In several cases, two equally significant coexistence intervals are obtained.

160 T. Utescher et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 253 (2007) 157–168

Author's personal copy

range of the palaeoclimate. For more details of theapproach the reader is referred to the above literature.

When using the CA, climate data of all knownNearest Living Relatives of the fossil taxa have to beextracted from the database PALAEOFLORA (Utescherand Mosbrugger, 1990–2007). To enable this process,floral lists first have to be checked with respect tosynonyms and a valid taxonomy, especially whenanalyzing floras described in older literature. Theselection of Nearest Living Relatives for each of thefossil taxa is then generated by the database and istherefore based on more recent palaeobotanical studies.In addition, fossil plant taxa with uncertain taxonomicposition as well as uncertain botanical affinity on thegeneric level are not considered when calculatingpalaeoclimate data.

In the present study, all floras are analyzed withrespect to 7 climate variables. These are mean annualtemperature (MAT), cold month mean (CMM), warmmonth mean WMM), mean annual precipitation (MAP),precipitation in the warmest month (MPwarm), precip-itation in the driest month (MPdry), and precipitation inthe wettest month (MPwet). The results obtained for thesingle climate variables are given in the Tables 3 and 4.

The number of taxa contributing with climate dataranges between 9 and 29 (mean: 20 taxa; cf. Table 3), and,thus, the diversity of the Nearest Living Relatives knownfor each flora is sufficient to produce reliable results whenapplying the CA (Mosbrugger and Utescher, 1997). Asstated above, taxonomically problematic forms and taxawith uncertain botanical affinity are excluded from theanalysis. The percentages of extant plant taxa that maycoexisting in the resulting climate intervals commonlyrange between 90% and 100% of all Nearest LivingRelatives known for the flora, or almost reach 90%,respectively (Tables 3 and 4). In all of these cases, theclimate data obtained from the analysis are considered ashighly significant (Mosbrugger and Utescher, 1997). Injust one case, a coexistence percentage below 85% results(Bela Stena,MAT; 83.3%; cf. Table 3) possibly caused bytaxonomic uncertainties. When applying the CA on theSerbian floral record no permanent climatic “outliers” areobserved. Permanent outliers in the CA analysis havebeen successfully used to detect misinterpretations of thepalaeobotanical record (Mosbrugger and Utescher, 1997).

3. Results

3.1. Outline of the floristic evolution

The oldest site analyzed in the present study is theearlier Oligocene Bogovina flora. With respect to its

floristic composition the flora is very specific andunusual and can be characterized as a subtropicalneedleleaved forest. Quasisequoia couttsiae and cryp-tomeroid type twigs with small tipped leaves (Taxodia-ceae vel Cupressaceae), formerly described as “Sequoia(Araucarites) sternbergii”, dominate the spectrum. Bothcomponents represent extinct conifer genera and arecommonly rare in the Serbian floral record. With respectto the edaphic conditions reported, the flora can beregarded as almost autochthonous. The presence ofAnthracotherium minus (Laskarev, 1925) most probablyconnects this flora with a final early Oligocene age andthe MP24 mammal zone (Pantic, 1984).

The Ravna Reka and Stamnica floras belong to thelatest Chattian (MP30 mammal zone) and the earlyMiocene, respectively. Both floras represent dominantlyevergreen vegetation types. In the Ravna Reka flora,legume-like taxa and other elements typical forxerothermic conditions are abundant, possibly pointingto a more pronounced seasonality of precipitation. In theStamnica flora, broadleaved evergreen genera, such asLaurus, Cinamomum (Daphnogene), Persea and Ficusdominate the spectrum. Deciduous floral elements arepoorly represented in both of the floras. At Ravna Reka,Alnus, Salix and Acer sporadically occur, while atStamnica, Castanea, Populus and Acer are recorded.The different aspect of both associations possibly can beattributed to microclimatic factors caused by theevolving palaeo-relief and/or different palaeoaltitude.In the lower Miocene, the relief of the northern part ofSerbia was only slightly raised, while in the southernpart, in the area of the present-day Dinaride Mountains,the relief was moderately raised, but no high mountainsexisted (Pantic and Dulic, 1992b).

The middle Miocene is represented by 8 localities.Popovac represents a typical Badenian flora, about time-equivalent with the Tuzla flora (Bosnia–Herzegovina)dated by marine fauna (Pantic, 1957), as well as by theoccurrence of mastodons and crocodiles (Lukovic, 1950;Pejovic, 1951; Petronijevic, 1967). Both floras existed onthe southern slopes of the Paratethys. The floral record ofPopovac comprises numerous laurophyllous elements,while deciduous taxa are sparse and possibly represent thevegetation of colline areas (e.g., Acer or Rhamnus). TheSlanci and Misaca floras also belong to the middleMiocene but are slightly younger than the Popovac flora.The latter is dated by marine fauna and is regarded as“fixed flora” (standard) for the younger parts of theBadenian (Pavlovic, 1903; Pantic, 1956). In the Slanciflora, members of the Lauraceae family are mostimportant (ca. 25 % of the taxa recorded), especiallyDaphnogene. In addition, other warm temperate

161T. Utescher et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 253 (2007) 157–168

Author's personal copy

elements, such as evergreen oaks and “Eucalyptus”,xeromorphic leaves of myrtaceous affinity, are recorded,as well as various legume-like taxa possibly representinga xerophilous, azonal vegetation type. Typical temperate,deciduous taxa, such as Salix, Alnus, Populus and Acer,but also Engelhardtia, Zelkova and conifers only reachminor percentages (ca. 15 % of the taxa recorded). Thehigh diversity of ferns observed at Slanci points to thepresence of rich undergrowth and more or less autoch-thonous conditions. In the Misaca flora, arcto-tertiaryfloristic elements are more important as is obvious fromthe presence of various species belonging to Acer,additionally, also Ulmus and Carpinus occur. These taxapossibly migrated to Serbia across the higher altitudes ofthe rising Carpathian mountain range. In the mixedmesophytic vegetation of Central Europe, deciduous,arcto-tertiary taxa played amore important role during theBadenian (e.g., Wieliczka–Viehhausen complex, cf. Mai,1995; Utescher et al., 2007-this volume).

Seliste, Bukovac (Saranovo), Bozdarevac, Bela Stenaand PancevoBridge are Sarmatian localities, and, in Serbia,there is a significantly large number of Sarmatian floraswith a rich palaeobotanical record. In the Seliste flora,typical temperate elements andwarm temperate taxa, partlywith a Mediterranean aspect, are about equally important.Among the 14 typical temperate genera reported, Liqui-dambar is most common. In addition, members of

Juglandaceae, as well as Populus, Salix, Zelkowa, andAlnus can be cited, obviously representing a riparian(gallery) forest association.Among the evergreen elements,Daphogene and Laurus are frequently present. Theoccurrence of Ziziphus, Nerium and legume-type taxa, incontrast, points to a certain seasonality of precipitation.While in Bukovac and Bozdarevac, there are againnumerous arcto-tertiary elements observed (Acer triloba-tum, Betula prisca, Alnus kefersteini, Fagus pliocenica,Carpinus grandis, Salix angusta, Salix longa, Salix media,Pupulus latior, Populus balsamoides, Ulmus longifolia,Ulmus carpinoides, Platanus aceroides), the situation isquite different in the slightly younger Bela Stena andPancevo Bridge floras. There, arcto-tertiary elements aresparse, while a larger number of taxa with lauraceousleaves, and partly also legume-like taxa are recorded. ThePancevo Bridge flora is dominated by taxa with laurel-likeleaves (55%) and conifers (32%). Ilex, Myrica, evergreenoaks and legume-type taxa are present reaching about 12%of the taxa recorded, while only 7% of taxa are typicaltemperate (e.g., Prunus, Rhamnus, Zelkowa, Juglans).Conifers, such as Sequoia, Libocedrus, andPinus also playan important role reaching about 32% of taxa. This leads tothe assumption that the floristic record represents a climaxforest composed of laurophyllous and conifer taxa, avegetation type presently common in higher altitudes undera warm temperate climate. The Bela Stena flora originates

Fig. 2. Coexistence intervals for mean annual temperature (MAT), cold month mean (CMM) and warm month mean (WMM). The arrows indicatestratigraphical uncertainties. The solid curves are tentatively plotted about connecting interval means. Dotted curves: climate record of the northwestGerman Cenozoic combined from published data (Utescher et al., 2000; Mosbrugger et al., 2005). The station data refer to the present-day climate.

162 T. Utescher et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 253 (2007) 157–168

Author's personal copy

from a sedimentary environment in which extremely small-grained, white, board-like marls were deposited. As in thePancevoBridge flora, high percentages of lauraceous formsand also many legume-like taxa are present, conifers, incontrast, are not important. The Bukovac flora represents acoastal vegetation type, while for the Bozdarevac andPancevo Bridge floras an island vegetation type with apronounced hilly-mountain relief is assumed (Pantic andDulic, 1992a). For the Bela Stena site, there is no certaininformation about the palaeo-relief.

The Late Miocene is represented by three localities.The Dubona flora belongs to the later Pannonian, theCrveni Breg and Osojna (Kladovo) floras to the upperpart of the Pontian. Arcto-tertiary elements by nowdominate the floral spectra although there is still asignificant number of subtropical species, some of themprobably had relict character even then. In the lateMiocene and the Pliocene, several regions of theMediterranean realm with obviously favourable micro-climatic conditions are known as a refuge area for so-called Central European Relicts (Kovar-Eder, 2003).Some of the Pontian sites in the study area representdiverse, coastal peatland vegetation. From these peatbogs large brown coal deposits developed in Serbia andin Bosnia–Herzegovina.

3.2. Palaeoclimate

The palaeoclimate data calculated by the coexistenceapproach (Tables 3 and 4) are shown in the Figs. 2 and 3where they are approximately plotted next to thechronological time-scale. In order to obtain climatecurves that can be easily compared with other recordsmeans of coexistence intervals are tentatively connectedinterpolating between the single floras. As is obviousfrom Fig. 2, highest temperatures result for the earlierOligocene Bogovina flora, with MATalmost up to 20 °Cand CMM around 10 °C. At the Oligocene/Miocenetransition, there is a pronounced temperature decreaseobserved. For the Ravna Reka and the Eggenburgian toOttnangian Stamnica flora a MAT of about 15 °C andCMM of about 4 °C is obtained. In the Badenian,temperatures rose again. CMM increased by about 4 °C(Popovac, Slanci, Misaca floras) while the temperatureexcursions observed for MAT and WMM are onlymodest. The warm climate phase in the Badeniancorresponds to a globally observed warm time interval,the Mid-Miocene Climate Optimum (Zachos et al.,2001). From the Sarmatian on, data indicate a more orless continuous cooling trend and a pronounced increasein seasonality of temperature. Until the Pontian, MAThas decreased by about 2.5 °C and CMM by 5 °C when

the means of coexistence intervals are regarded. WMMis not very much affected by this overall cooling trendstaying at the high level of about 26 °C. Among theSarmatian floras analyzed, Bela Stena and PancevoBridge tend to be warmer when compared to the onlyslightly older or even almost time-equivalent Bukovacand Bozdarevac floras. This tendency is most evidentfrom WMM values where a temperature difference ofalmost 2 °C can be stated.

As shown by the precipitation records (Figs. 2 and 3)annual precipitation rates above 800 mm are indicatedfor the time-span from the early Oligocene to the lateMiocene. Precipitation rates of the wettest month werewell above 100 mm and above 30 mm in the driestmonth, respectively, with the exception of the SarmatianBukovac flora where drier climate conditions may beassumed (cf. MPdry and MPwarm records on Fig. 3).For the early Oligocene Ravna Reka flora overall highvalues result for the different precipitation variables.During the prominent cooling in the later Oligocene (seeabove) MPwet dropped slightly (ca. 20 mm), while theother precipitation variables stayed about at the samelevel. For MAP, however, two coexistence intervals areobtained (Ravna Reka flora) with the drier one rangingbetween 897 and 975 mm (upper limit: Taxus) and thewetter between 1003 and 1375 mm (lower limit:Quercus cruciata with NLR Q. falcate) (Table 4).According to the method applied both intervals have thesame significance level and probability. Thus, it can beassumed that the flora comprises elements originatingfrom different stands with regionally higher and lowerrainfall, respectively. During the early Miocene, precip-itation rates increased again peaking in the Eggenbur-gian/Ottnangian (Popovac flora: MAP, MPwarm,MPwet). According to the present data the above time-interval represents the wettest period of the SerbianCenozoic. During the Middle Miocene, MPwet shows aclear decreasing trend while MPdry almost stays at thesame level. The MAP and MPwarm records, in contrast,are highly heterogeneous. Drier conditions result for theBadenian Slanci flora (MPwarm only), as well as for thelate Badenian/early Serravallian Bozdarevac and Pan-cevo Bridge floras while wetter conditions are indicatedfor the about contemporaneous Seliste, Bukovac andBela Stena floras. For the Misaca flora (MAP), as wellas for Bozdarevac (MPdry), there are again two equallysignificant coexistence intervals obtained with Laurussp. being the delimiting factor for the drier ranges andPersea sp. and Platanus occidentalis for the wetterintervals, respectively. During the late Miocene, there isa decreasing trend observed for MAP when consideringthe very high rates obtained for some of the late middle

163T. Utescher et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 253 (2007) 157–168

Author's personal copy

Miocene floras (Seliste, Bela Stena). The same is truefor MPwarm decreasing to ca. 80 mm in the Pontian andshifting in a direction to present day conditions(60 mm). MPdry stays at the same level and MPweteven shows a slight increasing trend.

4. Discussion

As shown by the present study, basic patterns of thepalaeoclimate evolution during the Cenozoic of Serbia

are well reflected by the vegetation changes observed.The high temperatures reported for the early Oligocenecorrespond to the absence of deciduous, arcto-tertiaryelements in the Bogovina flora. The steep gradient tolower temperatures during the late Oligocene present inthe Serbian record most probably is related to platetectonic movements and the northward shift of theAdriatic microplate at that time (Fig. 4; Popov et al.,2004). This cooling is also mirrored by vegetationchange. In the latest Oligocene and the early Miocene,

Fig. 3. Coexistence intervals for mean annual precipitation rates (MAP), precipitation in the warmest month (MPwarm), the driest month (MPdry),and the wettest month (MPwet). The solid curves are tentatively plotted about connecting interval means. Dotted curves: MAP record of the northwestGerman Cenozoic (Mosbrugger et al., 2005), records for other precipitation variables from Utescher et al. (2000) and new calculations. The stationdata refer to the present-day climate.

164 T. Utescher et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 253 (2007) 157–168

Author's personal copy

broadleaved evergreen taxa were still dominating thespectra, but by now, also several arcto-tertiary floristicelements were present in the Serbian floral recordmigrating to the Balkans from other parts of Europe(Ravna Reka, Stamnica). The subxeric aspect of theRavna Reka flora points to the fact that the coolingobserved during the late Oligocene obviously alsocaused drying, at least on a limited, regional scale. Thefact that for MAP rates partly two equally significantcoexistence intervals are obtained for a single florapoints to a small scale regional differentiation with drierand wetter stands represented by the palaeobotanicalrecord (Fig. 3). A subxeric aspect of plant associationsnear the Oligocene/Miocene boundary is also knownfrom other parts of the Central Paratethys, like from theSouth German Molasse Basin (Eger-Hausham floralcomplex) and the Swiss Molasse (Berger, 1990).

The warm climate phase observed during the middleMiocene (Fig. 2) coincides well with the dominantlyevergreen broadleaved vegetation with numerous laur-ophyllous taxa as reported for the Badenian Popovac,

Misaca and Slanci floras. As shown by studies on treediversity, around 80% of arboreal taxa described for thePopovac and Misaca floras are broadleaved evergreen, avalue among the highest of all other contemporaneousEuropean floras analyzed (Utescher et al., 2007-thisvolume). In most time-equivalent localities of theCentral Paratethys, in contrast, mixed mesophytic forestassociations existed. In these communities, evergreen,laurophyllous taxa and other elements known from thetypical Mastixoid floras also play an important role,however, deciduous taxa were dominating the spectra(e.g., Wieliczka–Viehhausen complex; Mai, 1995).

With the onset of the cooling in the Sarmatian,typical temperate, deciduous floristic elements becameincreasingly important (Seliste, Bukovac, Bozdarevac),and, also in Serbia the evergreen broadleaved vegetationwas replaced by a mixed mesophytic vegetation (e.g.,Utescher et al., 2007-this volume). In some places,however, broadleaved evergreen vegetation persisted(Bela Stena, Pancevo Bridge floras), obviously as arelict of the so-called Younger Mastixioid Floras widely

Fig. 4. Plaeogeographic maps showing the evolution of the Paratethys from the Oligocene to the Late Miocene (from Harzhauser and Piller, 2007-thisvolume). The arrow indicates the position of the study area.

165T. Utescher et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 253 (2007) 157–168

Author's personal copy

distributed in Central Europe during the early and theearly middle Miocene (e.g., Kovar-Eder, 2003). Forseveral of the Sarmatian floras, lower precipitation rateswith respect to different variables result from the applica-tion of the CA (Fig. 3; Table 4; e.g., MPdry/Seliste,MAP/Pancevo Bridge). In some cases, two coexistenceintervals are obtained, namely a “drier” and a “wetter”.In most of the cases, the presence a second, “drier” in-terval is correlated with higher diversities of sclero-phyllous taxa and/or taxa with legume-type leaves (e.g.,Seliste). According to most authors, these elements areindicative for subhumid conditions and seasonal drought.In the Central Paratethys realm, high diversities of theseplant types already occurred in earlier times, e.g., inKarpatian floras (Langenau–Leoben complex sensu Mai,1995) and in the Badenian (Utescher et al., 2007-thisvolume). In the Sarmatian, “dry floras” characterized bythe presence of numerous microphyllous taxa are mostcommon in an area stretching from the PannonianBasin to South Bohemia and eastern Austria while theclimatic conditions under which these associationsexisted are still controversially discussed (for a summarysee Mai, 1995). However, most of the authors reject thepossibility that in the Central Paratethys realm, a wide-spread xerothermic vegetation belt existed at that timebecause numerous taxa, present in the floral records,indicate overall humid climate conditions. Therefore,authors favour specific edaphic conditions restrictedto small-scale areas (e.g., Palamarev, 1989). In addition,it was shown that there is no good correlation betweenthe diversities of legume-like taxa and of sclerophyl-lous forms, both favoured as indicators for subhumidconditions, so far. According to the analyses of the ear-ly to middle Miocene floral record of Europe bothcomponents reach high percentages in almost differentregions (Jechorek and Kovar-Eder, 2004). This may leadto the assumption that the climatic interpretation ofthe legume-type leaves found in Miocene floras is stilldoubtful.

In the present approach, the palaeoclimatic interpre-tation of floras rich in legume-type leaves is certainlyincomplete. Since these forms cannot be assigned topresent taxa beyond the family level they do notcontribute with climate data to the analyses. However,the fact that, in the Sarmatian of Serbia, all over humidas well as seasonally dry vegetation types existed almostcontemporaneously, and the fact that “wet” and “dry”precipitation intervals may result for a single florasupport the theory of regionally differentiated precipi-tation patterns and special edaphic conditions, asproposed by Palamarev (1989). However, short-termchanges of precipitation rates in time cannot be excluded

so far because of the impreciseness of availablestratigraphical dating. According to the present data,precipitation rates in the Sarmatian were considerablyhigher than at present (MAP, MPwet) or at about anequal level (MPwarm, MPdry) even when the “dry”coexistence intervals are regarded. Under these con-strains no real xerophytic vegetation would have de-veloped at that time.

The dominance of deciduous taxa in the late Miocenefloras Dubona II, Crevni Breg and Osijina is obviouslyprimarily linked to the continuously decreasing CMM.On the other hand, climate can still be characterized aswarm temperate (Table 3) and favourable for warmthloving taxa. This coincides well with the presence of asignificant number of subtropical species and relicts, aspreviously mentioned, as well as the comparatively lateappearance of typical temperate taxa, such as Quercusrobur type in Serbia. From the composition of the lateMiocene floras analyzed there is no more evidence forthe existence of subhumid communities. Also in thePontian, vegetation has a humid aspect.

Although time resolution of the climate curvesobtained is limited, major trends, like the Mid-MioceneClimate Optimum, and the Late Miocene Cooling, arewell represented by the data. Thus, the continentalcurves can be correlated with global climate change asreflected in marine isotope records (e.g., Zachos et al.,2001). Additionally, correlations with other continentalclimate records are possible reconstructed for otherrealms of the European Cenozoic, e.g., the record for thenorthwest German Cenozoic (Utescher et al., 2000;Mosbrugger et al., 2005), for the Central Paratethys(Ivanov et al., 2002; Mosbrugger et al., 2005) andUkraine (Syabryaj et al., 2007-this volume). In Figs. 2and 3, the Serbian palaeoclimate record is plotted to-gether with curves for the northwest German Cenozoic(Lower Rhine Basin; Utescher et al., 2000; Mosbruggeret al., 2005). With the exception of the Oligocene datawhere plate tectonics might be the most important factorit is shown that temperatures generally are on a similarlevel. During the late Miocene, CMM decreased morepronounced and earlier in Serbia pointing to morecontinental type climate conditions. At present, theSerbian climate has significantly warmer summers whencompared to northwest Germany (Fig. 2). This wasobviously not true in the Cenozoic. At that time,extended water bodies (Paratethys, Lake Pannon) mighthave moderated the climate during the warm season.However, it is shown that temperatures in the lateMiocene of Serbia were only at an intermediate levelwhen compared to those calculated for other locations inthe mid-latitude Western Eurasia (Bruch et al., 2004,

166 T. Utescher et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 253 (2007) 157–168

Author's personal copy

2006). Thus, favourable temperatures were obviouslynot the most important factor for the forming of a refugearea in Serbia at that time.

Comparing the precipitation rates reconstructed for thenorthwest German Cenozoic to the Paratethys record,Serbian data tend to be lower in most of the floras anal-yzed, especially in the wettest and the warmest month.Only in parts of the Badenian, a similar high level asreconstructed for the Lower Rhine Basin is reached. Thiscoincides very well with the fact that in the Cenozoic,northwest Germany is part of the so-called CentralEuropean Wet Zone (Van Dam, 2006; Utescher et al.,2007-this volume) while Serbia has a marginal position toa somewhat drier continental area in the East. However, forthe major part of the time period regarded, the climate hasto be characterized as all over humid. Only for laterBadenian to Sarmatian floras there is stronger evidence forpartially drier conditions and possibly also for the pre-sence of a drier season. As obvious from the data calc-ulated forMPwarm andMPdry, however, it is shown thatthe driest season was not in summer, and, even then, noMediterranean type climate existed. Seasonally variableprecipitation rates without summer drought are alsoassumed for Sarmatian localities of the neighbouringPannonian area (e.g., Hungary; cf. Mai, 1995). Like inthe Messinian of the northern Mediterranean realm (e.g.,Bertini, 1994) a moist type of climate existed in Serbiaduring the Pontian.

5. Conclusions

We present the first detailed climate record for theSerbian Cenozoic which is discussed in the context ofvegetation change. The quantitative terrestrial climatedata can be correlated with global long-term trends andpatterns, such as the Mid-Miocene Climate Optimum andthe Late Miocene Cooling. On the other hand, regionaleffects, like moving tectonic plates and the presence ofwater bodies, are reflected by the climate data. As isobserved in the northwest German continental climaterecord the Late Miocene Cooling is most pronounced inwinter temperatures.

Basic patterns of the reconstructed climate evolutioncorrespond to fundamental changes in vegetation type.During the warm climate phase in the Badenian,dominantly evergreen vegetation existed. With the onsetof the cooling during the Sarmatian, it was replaced bymixed mesophytic forest and, later on, by dominantlydeciduous vegetation.

During most of the time period regarded mean annualprecipitation rates in Serbia were considerably higherthan at present. In the driest month, precipitation rates

were close present-day conditions. As is shown by theprecipitation data and by vegetational interpretationsthere is some evidence for seasonally dry conditions inthe Badenian and Sarmatian, at least on a regionalclimate scale. However, the problem of drier and wettervegetation associations co-occurring is far from beingcompletely understood, and further studies are required.In this context a taxonomic evaluation of the legume-like taxa present in the Serbian palaeobotanical recordand their quantitative climatic interpretation would beparticularly informative.

Acknowledgements

The authors like to thank all colleagues andNECLIME members for many fertile discussions. Weare indebted Sun Ge and J. Eder who carefully reviewedthe manuscript. This work would not have been possiblewithout the financial support granted by DFG. Thestudy presented herein is a contribution to the program“Neogene Climate Evolution in Eurasia— NECLIME”.

References

Berger, J.P., 1990. Le role des environnements de depot pour lesreconstitutions climatiques; les gisements a vegetaux de la molassegrise de Lausanne (Miocene inferieur, Suisse occidentale).Paleobiologie Continentale 17, 345–353.

Bertini, A., 1994. Palynological investigation on Upper Neogeneand Lower Pleistocene sections in Central and Northern Italy:Memorie de la Societa Geologica Italiana, 48, pp. 431–443.

Bruch, A.A., Utescher, T., Olivare, C.A., Dolakova, N., Ivanov, D.,Mosbrugger, V., 2004. Middle and Late Miocene spatial temperaturepatterns and gradients in Europe — preliminary results based onpalaeobotanical climate reconstructions. Courier ForschungsinstitutSenckenberg 249, 15–27.

Bruch,A.A.,Utescher, T.,Mosbrugger,V., Gabrielyan, I., Ivanov,D., 2006.Late Miocene climate in the circum-Alpine realm — a quantitativeanalysis of terrestrial palaeofloras. Palaeogeography, Palaeoclimatol-ogy, Palaeoecology 238, 270–280.

Harzhauser, M., Piller, W.E., 2007. Benchmark data of a changing sea-Palaeogeography, Palaeobiogeography and Events in the CentralParatethys during theMiocene. Palaeogeography, Palaeoclimatology,Palaeoecology 253, 8–31 (this volume) . doi:10.1016/j.palaeo.2007.03.031.

Ivanov, D., Ashraf, A.R., Mosbrugger, V., Palmarev, E., 2002.Palynological evidence for Miocene climate change in theForecarpathian Basin (Central Paratethys, NW Bulgaria). Palaeo-geography, Palaeoclimatology, Palaeoecology 178, 19–37.

Jechorek, H., Kovar-Eder, J., 2004. Vegetational characteristics in Europearound the late early to earlymiddleMiocene based on the plantmacrorecord. Courier Forschungsinstitut Senckenberg 249, 53–62.

Kovar-Eder, J., 2003. Vegetation dynamics in Europe during theNeogene. Deinsea 10, 373–392.

Kovar-Eder, J., Kvacek, Z., Zastawniak, E., Givulescu, R., Hably, L.,Mihajlovic, D., Teslenko, Y., Walther, H., 1996. Floristic trends inthe vegetation of the Paratethys surrounding areas during Neogene

167T. Utescher et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 253 (2007) 157–168

Author's personal copy

time. In: Bernor, R., Fahlbusch, V., Mittmann, H.-W. (Eds.), Theevolution of western Eurasian Neogene faunas. ColumbiaUniversity Press, pp. 399–409.

Laskarev, V., 1925. Sur la trouvaille des Anthracotherides en Serbie eten Bosnie. Annales Geologiques de la Peninsule Balkanique 8,32–50.

Lukovic, M.T., 1950. Geoloska ispitivanja u tercijaru okoline Popovcakod Paracina. Zbornik Radova Geoloskog Instituta SAN 3, 56–73.

Mai, D.H., 1995. Tertiäre Vegetationsgeschichte Europas. GustavFischer, Jena, Stuttgart, New York.

Mihajlovic, D., 1977. Neogene floras of “Balkan land” and theirsignificance for paleoclimatology, paleobiogeography and biostra-tigraphy III. Annales Geologiques de la Peninsule Balkanique 43-44,239–261.

Mihajlovic, D., 1978. Fossil flora from Slanci near Belgrade. Bulletindu Musee d'Historie Naturelle, serie A, livre 33.

Mihajlovic, D., Lazarevic, Z., 1999. Three late Pontian leaf floras fromnorthern Serbia reflecting different enviroments. Acta Palaeobo-tanica. Supplementum 2, 341–347.

Milovanovic, D., Mihajlovic, D., 1984. A Miocene flora from Zagubicabasin, eastern Serbia. Annales Geologiques de la PeninsuleBalkanique 48, 201–213.

Mosbrugger, V., Utescher, T., 1997. The coexistence approach — amethod for quantitative reconstructions of Tertiary terrestrialpalaeoclimate data using plant fossils. Palaeogeography, Palaeo-climatology, Palaeoecology 134, 61–86.

Mosbrugger, V., Utescher, T., Dilcher, D.L., 2005. Cenozoic continentalclimatic evolution of Central Europe. Proceedings of the NationalAcademy of Sciences 102 (42), 14964–14969.

Palamarev, E., 1989. Palaeobotanical evidence of the Tertiary historyand origin of the Mediterranean sclerophyll dendroflora. PlantSystematics and Evolution 162, 93–107.

Pantic, N.K., 1956. Biostratigraphie des flores tertiaires de Serbie.Annales Geologiques de la Peninsule Balkanique 24, 199–317.

Pantic, N.K., 1957. Biostratigraphie der tertiären Flora aus demsüdöstlichen Bosnien. Zbornik Drugi Kongres Geologa FNRJ,Sarajevo 185–195.

Pantic, N.K., 1984. On the evolution of land flora based on plant fossilsfrom the territory of Serbia. In: Jankovic, M. (Ed.), Vegetation de larepublique socialiste de Serbie. Academie serbie des sciences et desarts, Beograd, pp. 191–246.

Pantic, N.K., Dulic, I., 1992a. Study of lower Sarmatian fixed floras andtheir significance for reconstruction of land vegetation and phytos-tratigraphic correlations. Annales Geologiques de la PeninsuleBalkanique 56 (1), 105–118.

Pantic, N.K., Dulic, I., 1992b. On the regions of hill and the regions ofdominantly lowland vegetation during the early Miocene andgeodynamic events (the Dinarides — southern areas of Pannoniandepression). Annales Geologiques de la Peninsule Balkanique 56/2,83–100.

Pantic, N.K., Mihajlovic, D.S., 1977. Neogene floras of the Balkan landareas and their bearing on the study of palaeoclimatology, palaeo-biogeography and biostratigraphy (part 2). The lower Sarmatian floraof Beograd. Annales Geologiques de la Peninsule Balkanique 41.

Pantic, N.K., Mihajlovic, D.S., 1985. The problems of Eocene andOligocene in Serbia and further researching trends. GeoloskiGlasnik 28, 41–55.

Pavlovic, P., 1903. Prinove geoloskog zavoda II. Annales Geologiquesde la Peninsule Balkanique tome IV, 8–27 (in Serbian).

Pejovic, D., 1951. O nalasku vilice fosilnog krokodila u cementnimlaporcima Popovca kod Paracina. Zbornik radova SANU16, 103–107.

Petronijevic, Z.M., 1967. Die Mittelmiozäne und Untersarmatische(Steirische) Säugetierfauna Serbiens. Palaeontologia Jugoslavica7, 1–157.

Popov, S.V., Rögl, F., Rozanov, A.Y., Steininger, F.F., Shcherba, I.G.,Kovac, M., 2004. Lithological-Paleogeographic maps of Para-tethys; 10 maps Late Eocene to Pliocene. Courier Forschungsin-stitut Senckenberg 250, 46.

Rögl, F., 1998. Palaeogeographic considerations for Mediterraneanand Paratethys Seaways (Oligocene to Miocene). Annalen desNaturhistorischen Museums in Wien 99, 279–310.

Rögl, F., 1999. Mediterranean and Paratethys. Facts and hypotheses ofan Oligocene to Miocene paleogeography (short overview).Geologica Carpathica 50, 339–349.

Stevanovic, M.P., Pantic, N., 1954. O sarmatskoj flori i fauni izzeleznickih useka kod Bozdarevca. Annals of the GeologyPeninsuleBalkan XXII, 53–68.

Syabryaj, S., Utescher, T., Molchanov, S., 2007. Changes of climate andvegetation during the Miocene in the territory of Ukraine.Palaeogeography, Palaeoclimatology, Palaeoecology 253, 151–166(this volume) . doi:10.1016/j.palaeo.2007.03.038.

Utescher, T., Mosbrugger, V., 1990–2007. The Palaeoflora Database.(at http://www.palaeoflora.de).

Utescher, T., Mosbrugger, V., Ashraf, A.R., 2000. Terrestrial climateevolution in Northwest Germany over the last 25 million years.Palaios 15, 430–449.

Utescher, T., Erdei, B., Francois, L., Mosbrugger, V., 2007. Studies ondiversity of plant functional types in the Miocene of WesternEurasia- spatial distribution patterns in the Langhian, Sarmatian andTortonian, and their relation to palaeovegetation and palaeoclimate.Palaeogeography, Palaeoclimatology, Palaeoecology 253,224–248 (this volume) . doi:10.1016/j.palaeo.2007.03.041.

Van Dam, J.A., 2006. Geographic and temporal patterns in the lateNeogene (12-3 Ma) aridification of Europe. The use of smallmammals as paleoprecipitation proxies. Palaeogeography, Palaeo-climatology, Palaeoecology 238, 190–218.

Zachos, J.C., Pegani, M., Stone, L., Thomas, E., Billups, K., 2001.Trends, rhythms, and aberrations in global climates 65Ma to present.Science 292, 686–693.

All climate data provided here are accessible by the world database PANGAEA under the reference.Utescher, T., Djordjevic-Milutinovic, D., Bruch, A.A., Mosbrugger, V.,

2006. Oligocene to Miocene palaeoclimate reconstructions from 14sites in Serbia. PANGAEA. doi:10.1594/PANGAEA.472281.

168 T. Utescher et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 253 (2007) 157–168