Palaeogeography, Palaeoclimatology, Palaeoecology 304 (2011) 202–211

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Palaeogeography, Palaeoclimatology, Palaeoecology

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Precipitation patterns in the Miocene of Central Europe and the development of continentality

Angela A. Bruch a,⁎, Torsten Utescher b, Volker Mosbrugger a and NECLIME members 1 a Senckenberg Research Institute, Senckenberganlage 25, D-60325 Frankfurt a. M., Germany b Steinmann Institute, Bonn University, 53115 Bonn, Germany article info abstract

Article history: Understanding climate patterns, with their decisive influence on plant distribution and development, is Received 25 January 2010 crucial to understanding the history of vegetation patterns in Europe during the Miocene. This paper presents Received in revised form 8 October 2010 the detailed analyses of several precipitation parameters, including monthly precipitation of the wettest, Accepted 9 October 2010 driest and warmest months, for five Miocene stages. In conjunction with seasonality of temperature, those Available online 15 October 2010 parameters provide a meaningful measure of continentality and can help to document Miocene climate changes and patterns and their possible influence on vegetation. Climate reconstructions provided here are Keywords: entirely based on palaeobotanical material. In total, 169 Miocene floras were selected, including 14 Precipitation fl Continentality Burdigalian, 41 Langhian, 40 Serravallian, 36 Tortonian, and 38 Messinian localities. All oras were analysed Climate maps using the Coexistence Approach. The analysis of several precipitation parameters, the statistical inter- Europe correlation of results, and the comparison with modern patterns provides a comprehensive account on Open landscapes Miocene precipitation. Miocene climatic changes after the Mid Miocene Climatic Optimum (MMCO) are evidenced in our data set by three major factors, i.e. (1) increasing seasonality of temperature, (2) changes in the annual distribution of precipitation towards a precipitation peak in summer, and (3) a late increase of longitudinal gradients of precipitation parameters. Evidence of continental climate in Eastern Europe first appears during the Messinian. In addition to changes in temperature, shifts in the annual distribution of precipitation may have played a major role in post-Langhian climate changes. However, the most significant climatic transformations occurred later, from the end of Miocene through to the present. Several authors have described patterns of vegetation development in Europe that are in good agreement with our finding of the first evidence for continental climate in Eastern Europe during the Messinian. Our data do not support an onset of opening of vegetation during the Tortonian or even earlier, as has been described for some parts of Eastern and Southern Europe. Possibly either non climatic parameters influenced such an early development, or our data lack the required resolution and/or spatial coverage to fully decipher the influence of continentality on vegetation and to correlate climate and vegetation statistically. Nevertheless, climatic data that quantify continentality can provide a sound basis for explaining the expansion of grassland in Eurasia. © 2010 Elsevier B.V. All rights reserved.

1. Introduction climate today is typically found in conjunction with large areas of open landscape, with grassland predominant in the interior of Eurasia. Today the eastern and western coasts of Eurasia exist under very As part of the latter stage of the Cenozoic cooling, the Miocene was different climatic conditions influenced by the prevailing westerly a time of important climate and vegetation changes. In the Early atmospheric and oceanic circulation. The climate in western Eurasia is Miocene, glaciation was uni-polar, with an ice volume on Antarctica generally characterised by marine conditions along a west coast comparable to the present and a largely ice-free northern hemisphere. influenced by the Gulf Stream, with continentality increasing with During the Late Miocene, however, the first indications of northern distance from the coast. Climatologically, continentality is defined by hemispheric glaciation ultimately appeared leading to the formation a strong seasonality of temperature and low precipitation. Continental of the Greenland ice sheet in the Pliocene (Moran et al., 2006; Zachos et al., 2001). It is generally agreed that the epoch saw major environmental changes occurring both on the continents and in the ⁎ Corresponding author. Tel.: +49 69 7542 1568; fax: +49 69 746238. fi E-mail address: [email protected] (A.A. Bruch). oceans especially during the Late Miocene. A global intensi cation of 1 www.neclime.de orogenic movements considerably influenced the climate system; the

0031-0182/$ – see front matter © 2010 Elsevier B.V. All rights reserved. doi:10.1016/j.palaeo.2010.10.002 A.A. Bruch et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 304 (2011) 202–211 203 rapid uplift of the Tibetan Plateau, in particular, seems to have caused All floras have been analysed using the Coexistence Approach (CA) a stronger East Asian monsoon and triggered the upwelling systems following Mosbrugger and Utescher (1997). The method is one of the of the Indian Ocean (An et al., 2001). Likewise, the Late Miocene Nearest Living Relative Techniques that are based on the assumption witnessed the development and spread of C4-grasses, aridification of that the climatic requirements of Tertiary plant taxa are similar to the interiors of continents, and the expansion of open landscapes. those of their nearest living relatives (NLRs). With the CA, for each Although all these events are considered to be linked, there has as yet climate parameter the climatic ranges in which a maximum number been no proof of their causal interdependence (cf. Molnar, 2005). of NLRs of a given fossil flora can coexist is determined independently To understand the history of vegetation patterns in Europe, it is and considered the best description of the palaeoclimatic situation crucial to understand climate patterns as the main basis for plant under which the given fossil flora lived. distribution and development. Miocene temperature patterns have The application of the CA is facilitated by the computer program been discussed to date by Bruch et al. (2004, 2006, 2007) and by ClimStat and the database Palaeoflora which contains NLRs of more various authors in this issue (Liu et al., 2011-this issue; Utescher et al., than 3000 Cenozoic plant taxa, together with their climatic require- 2011-this issue; Yao et al., 2011-this issue). In the main, their data ments as derived from meteorological stations located within the demonstrate that the general cooling during the Miocene brought distribution areas of the taxa (see also information provided on the greater climatic differentiation, both spatially via increased latitudinal web site www.palaeoflora.de). According to the data available in the gradients and temporally via increased seasonality of temperature. In Palaeoflora data base, the method allows calculation of up to 15 addition to regional effects of palaeogeography such as the Paratethys climate parameters. sea and Alpine orogeny, temperature parameters reveal an increasing Typically, the resolution (width) and reliability of the resulting differentiation between marine and continental climate conditions. coexistence intervals increase with the number of taxa included in the However, that interpretation has largely been based on examinations analysis and are relatively high in floras with ten or more taxa for of temperature parameters. Miocene precipitation has been described which climate parameters are known. Because results of CA analyses so far only in terms of mean annual precipitation, whether through are intervals, the accuracy of calculated climate data corresponds to proxy-based reconstructions (e.g., Bruch et al., 2004, 2006, 2007; the accuracy of the borders of those coexistence intervals. Their Böhme et al., 2008, 2011-this issue; Mosbrugger et al., 2005; Utescher accuracy varies with respect to the parameter examined. It is highest et al., 2000) or in climate modelling (e.g., Micheels et al., 2007, 2009; for temperature-related parameters where it is usually within the Lunt et al., 2009). Studies of other precipitation parameters and the range of 1 to 2 °C and for mean annual precipitation with 100 to annual range of precipitation are lacking. Mertz-Kraus et al. (2009) 200 mm. Other precipitation parameters are less accurate and mainly interpret changes in coral growth increments as a signal of increased reflect overall trends. Although Mosbrugger and Utescher (1997) do winter rain in Crete at 9 Ma and as first evidence for Mediterranean- not give error bars for CA results of different parameters, their type climate. For the Pannonian basin, Harzhauser et al. (2007) application of the method to modern floras led them to conclude that postulate a peak in summer precipitation in the Late Miocene (ca. reconstructions of mean annual precipitation and of precipitation of 10 Ma) based on isotope data. Only van Dam (2006) provides a more the warmest month are least reliable and therefore not easy to exhaustive overview of changes and patterns of driest month precip- interpret. Thus, a correlation analysis (Section 3.1) is applied to itation from 12 to 3 Ma in Europe, basing the analysis on small mammal overcome these difficulties. For a detailed discussion and introduction communities. to the method, see Mosbrugger and Utescher (1997), Mosbrugger This paper will therefore focus mainly on a detailed analysis of (1999), and Utescher et al. (2000). precipitation parameters, including monthly precipitation of the In this study, mean annual precipitation (MAP), monthly precip- wettest, driest and warmest months. In combination with seasonality itation of the driest month (LMP), monthly precipitation of the of temperature, they constitute a reliable measure of continentality wettest month (HMP), and monthly precipitation of the warmest and can help document changes and patterns of Miocene climate and month (WMP) have been calculated according to the CA. In addition, their possible influence on vegetation. the mean annual range of temperature (MART — the temperature difference between of warmest and coldest months) and mean annual 2. Material and methods range of precipitation (MARP — the difference between HMP and LMP) have been taken into consideration. These parameters provide Climate reconstructions provided here are based entirely on detailed information on precipitation patterns and a means of as- palaeobotanical material. In total, 169 Miocene floras have been sessing continentality. selected, including 14 Burdigalian (20.428–16.303 Ma), 41 Langhian As stated above, the CA calculates for all climate parameter (16.303–13.654 Ma), 40 Serravallian (13.654–11.600 Ma), 36 Torto- coexistence intervals that are assumed to encompass the “real climate nian (11.600–7.251 Ma), and 38 Messinian (7.251–5.332 Ma) local- value”. For the purpose of data visualisation, the middle values of the ities, excluding the Aquitanian stage due to low data coverage calculated coexistence intervals are used. An analysis of recent (absolute ages after Harzhauser and Piller, 2007). Except for the European climate by Klotz (1999); see also (Pross et al., 2000) Burdigalian with only 14 samples, all other stages are represented by shows for subtropical to temperate conditions that the centres of the comparable numbers of floras. The selection attempts to provide coexistence intervals correlate better to the real data than do the sufficient data coverage of the area of interest on the one hand and to borders of the intervals. The use of middle values turns out to be narrow the stratigraphic range as much as possible on the other. To appropriate in visualising climatic trends between regions, especially increase the reliability of results, low-diversity floras were avoided. when interpolating over greater distances. Most of the investigated floras have been published already Maps of the reconstructed climate data were generated using the through the NECLIME network and are available in the PANGAEA GIS program ArcView. Interpolations between data points were database (www.pangaea.de; Bruch et al., 2004, 2006, 2007). New data calculated using the inverse distance weighted method, which will be published in PANGAEA with their geographic and stratigraphic provides a relatively smooth gradient between individual data points positions and references to the original palaeobotanical investigations. and more detailed patterns between neighbouring points, with less The complete data are also appended to this paper as Supplements 1 detail between points separated by greater distances. The interpola- and 2. All floras are from either continental or Paratethyan sediments; tion does not take into account altitudinal factors such as lapse rates. that restriction is intended to ensure relatively low variation in For areas without data, the program extrapolates climate patterns. To taphonomic influences (e.g., exclusion of long-distance transport into discourage overinterpretation of the resulting maps, we clearly marine sediments). identify within these maps the underlying data points (localities). 204 A.A. 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The maps are intended merely to aid understanding of the data set. various degrees but always in a similar way. When the different time Such caveats apply all the more to the maps that show the difference intervals are compared, the correlation coefficients between HMP and between fossil and modern data; their ostensible high resolution is WMP are seen to increase continuously from the Middle to the Late owed purely to the detailed modern WORLDCLIM data set (available Miocene. Thus WMP, as estimated using CA, is not only a good mea- at www.worldclim.org). Despite those reservations, such maps are sure for summer precipitation but can be assumed to reflect general valid visualisation tools with high value to assist the comparison trends and patterns in precipitation parameters, especially for post- among climatic states at different points in time. Langhian times. The graphical regressions shown in Fig. 1 indicate that the gradient 3. Results of regression between WMP and the other precipitation parameters is steepest in Burdigalian, Serravallian and Messinian data sets. The low 3.1. Correlation of precipitation parameters number of data available for the Burdigalian stage does not allow for further interpretation, but the generally increasing gradient in post- To facilitate the study of Miocene precipitation patterns in Europe, Langhian data evidences an increase in seasonal differentiation of mean annual precipitation (MAP), monthly precipitation of the driest precipitation towards a more pronounced precipitation peak in month (LMP), monthly precipitation of the wettest month (HMP), summer. On the other hand, despite high absolute values for summer monthly precipitation of the warmest month (WMP), and the mean precipitation and a high correlation coefficient, Langhian data show annual range of precipitation (MARP — the difference between HMP the lowest regressional gradient, reflecting generally humid condi- and LMP) were estimated via CA. In the first step, the entire data set tions with no specific wetter or drier season in summer. underwent a correlation analysis to determine the most significant parameters and the role of the wettest month data (tab. 1 and Fig. 1). Correlation coefficient and gradient of regression are two measures 3.2. Patterns of continentality that reflect the relationships between two parameters. Where the correlation coefficient expresses how reliably the data are related to Continentality is defined by low precipitation and strong season- one another, the regressional gradients show the nature of those ality of temperature. Figs. 2 and 3 reflect the history of continental relationships. climate in central Europe by showing the patterns of MART and MAP, The strongest correlation with the highest correlation coefficients both as palaeo-data alone and as differences of palaeo-minus-modern (N0.9, Table 1) in the entire data set arises between HMP and WMP. data. Fig. 4 provides patterns of WMP that indicate the course of the Both parameters correlate with the other precipitation parameters, to development of summer rain.

250 300

225 275

200 250

175 225

150 200 HMP [mm]

MARP [mm] 125 175

100 150

75 125

50 100 50 100 150 50 100 150 WMP [mm] WMP [mm]

1,600 60

1,400 50

1,200 40

30 1,000 LMP [mm] MAP [mm] 20 800

10 600

0 50 100 150 50 100 150 WMP [mm] WMP [mm]

Fig. 1. Correlation between WMP and other precipitation parameters; red circle — Burdigalian, blue triangle — Langhian; green reversed triangle — Serravallian; purple square — Tortonian; yellow rhombus — Messinian. A.A. Bruch et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 304 (2011) 202–211 205

Table 1 The differences between modern and palaeo-data document the Correlation coefficients between all precipitation parameters (Pearsons r) for each very low degree of change in precipitation (MAP, WMP) and MART Miocene stage; most significant values N0.9 are given in bold. during the Miocene compared to the transformations that occurred MAP HMP LMP WMP MARP post-Miocene and up to the present. Although absolute values are Messinian (n=38) clearly changing with the approach of the Messinian, they are still HMP 0.403021 remote from the modern situation. LMP 0.531935 0.696901 WMP 0.303088 0.982339 0.650542 MARP 0.212944 −0.521682 −0.208308 −0.672101 3.3. Seasonality gradients MART −0.140436 −0.440914 −0.153133 −0.479250 0.440318 To aid in analysis of the latitudinal gradients of precipitation and Tortonian (n=36) HMP 0.608278 MART as measures of continentality, fossil climate data are plotted LMP 0.315546 0.740779 against longitude in Fig. 5, while correlation coefficients are given in WMP 0.520197 0.979634 0.736951 Table 2. Here none of the data reveal statistically significant MARP 0.039588 −0.478946 −0.404016 −0.645457 longitudinal gradients. Only Messinian and Burdigalian data (with a MART −0.279421 −0.672349 −0.471745 −0.742259 0.687520 small data sample) show a correlation between longitude and WMP, Serravallian (n=40) and Messinian and Serravallian data to some extent with MAP. HMP 0.551581 The lack of significant correlation between longitude and MART LMP 0.571816 0.641429 can mainly be traced to the aforementioned buffering effect of the WMP 0.513372 0.973449 0.594771 Paratethys, which suppressed any development towards temperature MARP −0.031048 −0.237586 −0.026666 −0.453626 MART −0.336340 −0.493868 −0.481734 −0.465477 0.052502 seasonality in the Pannonian Basin. Nevertheless, data from the Messinian stage show a slight, although not statistically significant, Langhian (n=41) increase in MART, together with the strongest decrease in MAP and HMP 0.305209 WMP from west to east of all data. With its combination of LMP 0.361196 0.441367 characteristics, the Messinian stands alone as the only stage to display WMP 0.315189 0.952162 0.390299 MARP −0.175075 −0.324336 −0.050431 −0.597894 a tendency towards higher continentality in Eastern Europe. All other MART −0.026423 −0.155001 0.130017 −0.238667 0.332205 Miocene stages show either no longitudinal changes, or not in this combination. In short, evidence for continental climate in Eastern Burdigalian (n=14) Europe first appears with the Messinian. HMP 0.557893 LMP 0.381232 0.699585 WMP 0.536028 0.970741 0.664921 4. Discussion MARP −0.308981 −0.587317 −0.363037 −0.764483 MART −0.533929 −0.793751 −0.712488 −0.735265 0.347334 4.1. Miocene climate All data HMP 0.408527 To date, interpretation of Miocene climate has mainly been based LMP 0.041926 −0.465211 WMP 0.483416 0.605831 −0.177360 on temperature parameters, with Miocene rainfall generally described MARP 0.336634 0.972750 −0.657772 0.561957 in terms of mean annual precipitation. The current study relies on a MART −0.248167 −0.509534 0.376803 −0.379528 −0.532256 more comprehensive set of Miocene precipitation data and employed more precise precipitation parameters. On the whole, the data document generally humid conditions, with all parameters exceeding The mean annual range of temperature (MART, Fig. 2) is lowest present values. General climatic development over time can be sum- during Burdigalian and Langhian in Western Europe and increases marised as follows. beginning with the Middle Miocene. However, the coastal parts of the Langhian data show very high humidity, with a peak in pre- continent display only minor changes, reflecting a persistent low cipitation that does not match summer precipitation and therefore seasonality of temperature influenced by the Atlantic Ocean. This occurred other than in summer, and no longitudinal gradient. pattern is the first evidence for a Neogene climatic differentiation Serravallian data show slightly less humid conditions, evidence of between oceanic and continental climate in Europe. Due to the buf- a summer peak in precipitation, and no significant longitudinal fering effect of the large Paratethys sea, temperatures in Eastern gradient. Europe also remain equable at least until the Tortonian (Bruch et al., Tortonian data evidence slightly higher humidity than in previous 2004, 2006, 2007). During the Miocene, MART patterns were evi- times with increasing evidence of a summer peak in precipitation, but dently not as strictly east–west oriented as they are today. no longitudinal gradients. Miocene precipitation data all show generally humid conditions Messinian data evidence less humid conditions, increasing for the entire Miocene. Besides a general decrease in mean annual evidence of a summer peak in precipitation, and first evidence of precipitation (MAP, Fig. 3), the Langhian and Tortonian appear to be longitudinal gradients in WMP and MAP. wetter phases compared to the times before and after. Moreover, a This succession testifies to a general decrease in precipitation since progressive spatial differentiation appears in the late Middle Miocene the Middle Miocene and the onset of continental climate conditions in (Serravallian) with lower precipitation in the eastern part of central Eastern Europe in the Late Miocene. Europe. Beyond the general decrease in MAP, the Langhian and Tortonian Summer precipitation (WMP, Fig. 4) decreases beginning with the appear to have been wetter than the stages before and after. Wetter Middle Miocene, and all stages to some extent show a longitudinal conditions during the Langhian may be related to the Mid Miocene differentiation, with slightly lower values in eastern Europe. This Climatic Optimum (MMCO) and are widely discussed in the literature trend becomes more obvious towards the Late Miocene and is the (e.g., Kürschner et al., 2008; Retallack, 2009; Wan et al., 2009; You most prominent observed pattern change of all precipitation para- et al., 2009). It is worth noting for general discussion of MMCO that meters studied. Beyond changes in temperature, shifts in the annual our data show a strong annual range of precipitation in central Europe distribution of precipitation may have played a major role in post- together with the highest values for all precipitation parameters, Langhian climate changes. reflecting generally very humid conditions with some seasonality, but 206 A.A. Bruch et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 304 (2011) 202–211

Fig. 2. Mean annual range of temperature. Left column: absolute proxy data, IDW interpolated; green — high seasonality, red — low seasonality. Right column: Differences between interpolated reconstructions and modern WORLDCLIM raster data: green — past higher than present, white — past similar to present, red — past lower than present (for detailed legend see Fig. 4b). without a precipitation peak during summer (see also Böhme et al., mainly because the focus so far has been on comparing Middle and 2007). Late Miocene conditions and on the general decrease in humidity. The prevalence of wetter conditions during the Tortonian than in Only Böhme et al. (2008, and 2011-this issue) describe wetter preceding and successive stages has been less thoroughly discussed, conditions during the Tortonian than in the late Serravallian and early A.A. Bruch et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 304 (2011) 202–211 207

Fig. 3. Mean annual precipitation. Left column: absolute proxy data, IDW interpolated; blue — high precipitation, yellow — low precipitation. Right column: Differences between interpolated reconstruction and modern WORLDCLIM raster data: blue — past higher than present, white — past similar to present, brown — past lower than present (for detailed legend see Fig. 4b). All Miocene values are much higher than modern annual precipitation data.

Messinian, based on a study of herpetofauna and in good agreement confirm such low values for any of the parameters analysed. This may with our results. On the other hand, their data indicate lower than reflect the challenge of reconstructing very dry conditions on the basis present MAP during short-term dryer phases, where our data do not of plant fossils (see Böhme et al., 2006, 2007). The lack of fossil floras 208 A.A. Bruch et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 304 (2011) 202–211 preserved under dry conditions causes a strong bias in our data set with Moreover, in central and eastern Europe as well, faunas and floras gaps towards southern Europe, where plant proxy data are available usually come from different stratigraphic levels and taphonomic only from moister regions of Spain (NW coast) and Italy (N Italy). settings. It may well be that our data lack the necessary temporal

Fig. 4. Precipitation of the warmest month. Left column: absolute proxy data, IDW interpolated; blue — high precipitation, yellow — low precipitation. Right column: Differences between interpolated reconstruction and modern WORLDCLIM raster data: blue — past higher than present, white — past similar to present, brown — past lower than present (for detailed legend see Fig. 4b). All Miocene values are much higher than modern summer precipitation data. b. Legend of Figs. 2–4. A.A. Bruch et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 304 (2011) 202–211 209

Fig. 4 (continued). resolution and taphonomic ability to detect dry events that would support the results of Böhme et al. (2008, and 2011-this issue). However, aside from the probable bias between faunal and floral proxy data, the same basic signals may be documented by both studies. Not only the more humid Tortonian stage, but also an increasing aridification prograding from eastern Europe as well, as postulated by Böhme et al. (2011-this issue),isconfirmed by our data. That signal is revealed in our data with especial clarity in the increasing longitudinal Fig. 5. Longitudinal gradients of MART, MAP, and WMP; red circle — Burdigalian, blue gradients of MAP and WMP over time (Fig. 5). triangle — Langhian; green reversed triangle — Serravallian; purple square — Tortonian; — fi Miocene climatic changes after MMCO are testified in our data yellow rhombus Messinian. Correlation coef cients in Table 2. set by three major factors: (1) increasing seasonality of tempera- ture, (2) changes in the annual distribution of precipitation towards aprecipitationpeakinsummer,and(3)alateincreaseoflongi- tudinal gradients of precipitation parameters. addition, Syabryaj et al. (2007) describe the vegetation development of the Ukrainian plain as a stepwise opening of the forests, with the 4.2. Landscape opening in Europe first steppe grasslands arising during the early Tortonian (Kherso- nian) and expanding considerably during late Messinian (Pontian). The expansion of open landscapes during the Miocene in Europe The same pattern of development with an onset of opening of has been the subject of widespread and sometimes heated debate vegetation at the Khersonian stage is observed by Shatilova et al. based on fossil fauna and flora. Strömberg et al. (2007) provide a (2010) for a shallow-marine succession in eastern Georgia. Only comprehensive review of the discussion. Large mammal data speak Kovar-Eder et al. (2008) provide a broader view of the European for the presence of open environments in southern Europe since the vegetation history, based on a quantitative analysis of a huge floristic early Late Miocene (e.g., Agustí et al., 1999; Fortelius et al., 2006). data set from Europe. The authors summarise that for the Tortonian, However, those data mainly confirm mosaic landscapes with open forests and do not support the notion of vast open landscapes (van Dam and Reichert, 2009; and references herein). With regard to plant Table 2 fossils, only some data on phytoliths and pollen favour grass- Correlation coefficients between longitude and climate parameters (Pearsons r) for dominated savannas or open woodlands. Showing that such habitats each Miocene stage. were widely established in the eastern parts of southern Europe MART MAP WMP during the Late Miocene based on phytolith analyses, Strömberg et al. Messinian (n=38) 0.2595 −0.3517 −0.492 (2007) propose that relatively open habitats developed in Asia Minor Tortonian (n=36) −0.0862 0.2434 0.1649 beginning already in the Early Miocene. For Akgün et al. (2007), Serravallian (n=41) 0.1782 −0.4078 −0.1312 however, analysis of Anatolian pollen profiles documents increasing Langhian (n=40) 0.2717 −0.1932 −0.0154 Burdigalian (n=14) 0.5436 0.0551 −0.4388 abundance of open vegetation taxa only during the Tortonian. In 210 A.A. Bruch et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 304 (2011) 202–211

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Our data cannot be correlated with an onset of opening climate in the circum-Alpine realm — a quantitative analysis of terrestrial fl – of vegetation during or prior to the Tortonian. Either that develop- palaeo oras. Palaeogeography, Palaeoclimatology, Palaeoecology 238, 270 280. Bruch, A.A., Uhl, D., Mosbrugger, V., 2007. Miocene Climate in Europe — Patterns and ment in vegetation is not related to the climate parameters analysed Evolution. — a first synthesis of NECLIME. Palaeogeography, Palaeoclimatology here, or, conceivably other maybe non climatic parameters played a Palaeoecology 253, 1–7. — role, as discussed controversially by various authors for example for Cowling, S.A., 1999. Plants and temperature CO2 uncoupling. Science 285 (5433), 1500–1501. atmospheric CO2 concentration (e.g., Pagani et al., 1999; Cowling, Fortelius, M., Eronen, J., Liu, L., Pushkina, D., Tesakov, A., Vislobokova, I., Zhang, Z., 2006. 1999; Kürschner et al., 2008) or for the coevolution of grasses and Late Miocene and Pliocene large land mammals and climatic changes in Eurasia. grazers (Retallack, 2001). It is also possible that the implications of Palaeogeography, Palaeoclimatology, Palaeoecology 238, 219–227. Harzhauser, M., Piller, W.E., 2007. Benchmark data of a changing sea—palaeogeography, certain taxa as evidence for dryness and openness (e.g., Poaceae and palaeobiogeography and events in the Central Paratethys during the Miocene. Asteraceae) are overestimated by qualitative approaches or under- Palaeogeography, Palaeoclimatology, Palaeoecology 253, 8–31. estimated by our method due to the usually wide climatic ranges of Harzhauser, M., Latal, C., Piller, W.E., 2007. 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