Palaeogeography, Palaeoclimatology, Palaeoecology 304 (2011) 202–211
Contents lists available at ScienceDirect
Palaeogeography, Palaeoclimatology, Palaeoecology
j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / p a l a e o
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
b
Senckenberg Research Institute, Senckenberganlage 25, D-60325 Frankfurt a. M., Germany
Steinmann Institute, Bonn University, 53115 Bonn, Germany
a r t i c l e
i n f o
Article history:
Received 25 January 2010
Received in revised form 8 October 2010
Accepted 9 October 2010
Available online 15 October 2010
Keywords:
Precipitation
Continentality
Climate maps
Europe
Open landscapes
a b s t r a c t
Understanding climate patterns, with their decisive influence on plant distribution and development, is
crucial to understanding the history of vegetation patterns in Europe during the Miocene. This paper presents
the detailed analyses of several precipitation parameters, including monthly precipitation of the wettest,
driest and warmest months, for five Miocene stages. In conjunction with seasonality of temperature, those
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
entirely based on palaeobotanical material. In total, 169 Miocene floras were selected, including 14
Burdigalian, 41 Langhian, 40 Serravallian, 36 Tortonian, and 38 Messinian localities. All floras were analysed
using the Coexistence Approach. The analysis of several precipitation parameters, the statistical intercorrelation of results, and the comparison with modern patterns provides a comprehensive account on
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
Today the eastern and western coasts of Eurasia exist under very
different climatic conditions influenced by the prevailing westerly
atmospheric and oceanic circulation. The climate in western Eurasia is
generally characterised by marine conditions along a west coast
influenced by the Gulf Stream, with continentality increasing with
distance from the coast. Climatologically, continentality is defined by
a strong seasonality of temperature and low precipitation. Continental
⁎ Corresponding author. Tel.: + 49 69 7542 1568; fax: + 49 69 746238.
E-mail address: [email protected] (A.A. Bruch).
1
www.neclime.de
0031-0182/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2010.10.002
climate today is typically found in conjunction with large areas of
open landscape, with grassland predominant in the interior of Eurasia.
As part of the latter stage of the Cenozoic cooling, the Miocene was
a time of important climate and vegetation changes. In the Early
Miocene, glaciation was uni-polar, with an ice volume on Antarctica
comparable to the present and a largely ice-free northern hemisphere.
During the Late Miocene, however, the first indications of northern
hemispheric glaciation ultimately appeared leading to the formation
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
oceans especially during the Late Miocene. A global intensification of
orogenic movements considerably influenced the climate system; the
A.A. Bruch et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 304 (2011) 202–211
rapid uplift of the Tibetan Plateau, in particular, seems to have caused
a stronger East Asian monsoon and triggered the upwelling systems
of the Indian Ocean (An et al., 2001). Likewise, the Late Miocene
witnessed the development and spread of C4-grasses, aridification of
the interiors of continents, and the expansion of open landscapes.
Although all these events are considered to be linked, there has as yet
been no proof of their causal interdependence (cf. Molnar, 2005).
To understand the history of vegetation patterns in Europe, it is
crucial to understand climate patterns as the main basis for plant
distribution and development. Miocene temperature patterns have
been discussed to date by Bruch et al. (2004, 2006, 2007) and by
various authors in this issue (Liu et al., 2011-this issue; Utescher et al.,
2011-this issue; Yao et al., 2011-this issue). In the main, their data
demonstrate that the general cooling during the Miocene brought
greater climatic differentiation, both spatially via increased latitudinal
gradients and temporally via increased seasonality of temperature. In
addition to regional effects of palaeogeography such as the Paratethys
sea and Alpine orogeny, temperature parameters reveal an increasing
differentiation between marine and continental climate conditions.
However, that interpretation has largely been based on examinations
of temperature parameters. Miocene precipitation has been described
so far only in terms of mean annual precipitation, whether through
proxy-based reconstructions (e.g., Bruch et al., 2004, 2006, 2007;
Böhme et al., 2008, 2011-this issue; Mosbrugger et al., 2005; Utescher
et al., 2000) or in climate modelling (e.g., Micheels et al., 2007, 2009;
Lunt et al., 2009). Studies of other precipitation parameters and the
annual range of precipitation are lacking. Mertz-Kraus et al. (2009)
interpret changes in coral growth increments as a signal of increased
winter rain in Crete at 9 Ma and as first evidence for Mediterraneantype climate. For the Pannonian basin, Harzhauser et al. (2007)
postulate a peak in summer precipitation in the Late Miocene (ca.
10 Ma) based on isotope data. Only van Dam (2006) provides a more
exhaustive overview of changes and patterns of driest month precipitation from 12 to 3 Ma in Europe, basing the analysis on small mammal
communities.
This paper will therefore focus mainly on a detailed analysis of
precipitation parameters, including monthly precipitation of the
wettest, driest and warmest months. In combination with seasonality
of temperature, they constitute a reliable measure of continentality
and can help document changes and patterns of Miocene climate and
their possible influence on vegetation.
2. Material and methods
Climate reconstructions provided here are based entirely on
palaeobotanical material. In total, 169 Miocene floras have been
selected, including 14 Burdigalian (20.428–16.303 Ma), 41 Langhian
(16.303–13.654 Ma), 40 Serravallian (13.654–11.600 Ma), 36 Tortonian (11.600–7.251 Ma), and 38 Messinian (7.251–5.332 Ma) localities, excluding the Aquitanian stage due to low data coverage
(absolute ages after Harzhauser and Piller, 2007). Except for the
Burdigalian with only 14 samples, all other stages are represented by
comparable numbers of floras. The selection attempts to provide
sufficient data coverage of the area of interest on the one hand and to
narrow the stratigraphic range as much as possible on the other. To
increase the reliability of results, low-diversity floras were avoided.
Most of the investigated floras have been published already
through the NECLIME network and are available in the PANGAEA
database (www.pangaea.de; Bruch et al., 2004, 2006, 2007). New data
will be published in PANGAEA with their geographic and stratigraphic
positions and references to the original palaeobotanical investigations.
The complete data are also appended to this paper as Supplements 1
and 2. All floras are from either continental or Paratethyan sediments;
that restriction is intended to ensure relatively low variation in
taphonomic influences (e.g., exclusion of long-distance transport into
marine sediments).
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All floras have been analysed using the Coexistence Approach (CA)
following Mosbrugger and Utescher (1997). The method is one of the
Nearest Living Relative Techniques that are based on the assumption
that the climatic requirements of Tertiary plant taxa are similar to
those of their nearest living relatives (NLRs). With the CA, for each
climate parameter the climatic ranges in which a maximum number
of NLRs of a given fossil flora can coexist is determined independently
and considered the best description of the palaeoclimatic situation
under which the given fossil flora lived.
The application of the CA is facilitated by the computer program
ClimStat and the database Palaeoflora which contains NLRs of more
than 3000 Cenozoic plant taxa, together with their climatic requirements as derived from meteorological stations located within the
distribution areas of the taxa (see also information provided on the
web site www.palaeoflora.de). According to the data available in the
Palaeoflora data base, the method allows calculation of up to 15
climate parameters.
Typically, the resolution (width) and reliability of the resulting
coexistence intervals increase with the number of taxa included in the
analysis and are relatively high in floras with ten or more taxa for
which climate parameters are known. Because results of CA analyses
are intervals, the accuracy of calculated climate data corresponds to
the accuracy of the borders of those coexistence intervals. Their
accuracy varies with respect to the parameter examined. It is highest
for temperature-related parameters where it is usually within the
range of 1 to 2 °C and for mean annual precipitation with 100 to
200 mm. Other precipitation parameters are less accurate and mainly
reflect overall trends. Although Mosbrugger and Utescher (1997) do
not give error bars for CA results of different parameters, their
application of the method to modern floras led them to conclude that
reconstructions of mean annual precipitation and of precipitation of
the warmest month are least reliable and therefore not easy to
interpret. Thus, a correlation analysis (Section 3.1) is applied to
overcome these difficulties. For a detailed discussion and introduction
to the method, see Mosbrugger and Utescher (1997), Mosbrugger
(1999), and Utescher et al. (2000).
In this study, mean annual precipitation (MAP), monthly precipitation of the driest month (LMP), monthly precipitation of the
wettest month (HMP), and monthly precipitation of the warmest
month (WMP) have been calculated according to the CA. In addition,
the mean annual range of temperature (MART — the temperature
difference between of warmest and coldest months) and mean annual
range of precipitation (MARP — the difference between HMP and
LMP) have been taken into consideration. These parameters provide
detailed information on precipitation patterns and a means of assessing continentality.
As stated above, the CA calculates for all climate parameter
coexistence intervals that are assumed to encompass the “real climate
value”. For the purpose of data visualisation, the middle values of the
calculated coexistence intervals are used. An analysis of recent
European climate by Klotz (1999); see also (Pross et al., 2000)
shows for subtropical to temperate conditions that the centres of the
coexistence intervals correlate better to the real data than do the
borders of the intervals. The use of middle values turns out to be
appropriate in visualising climatic trends between regions, especially
when interpolating over greater distances.
Maps of the reconstructed climate data were generated using the
GIS program ArcView. Interpolations between data points were
calculated using the inverse distance weighted method, which
provides a relatively smooth gradient between individual data points
and more detailed patterns between neighbouring points, with less
detail between points separated by greater distances. The interpolation does not take into account altitudinal factors such as lapse rates.
For areas without data, the program extrapolates climate patterns. To
discourage overinterpretation of the resulting maps, we clearly
identify within these maps the underlying data points (localities).
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The maps are intended merely to aid understanding of the data set.
Such caveats apply all the more to the maps that show the difference
between fossil and modern data; their ostensible high resolution is
owed purely to the detailed modern WORLDCLIM data set (available
at www.worldclim.org). Despite those reservations, such maps are
valid visualisation tools with high value to assist the comparison
among climatic states at different points in time.
3. Results
3.1. Correlation of precipitation parameters
3.2. Patterns of continentality
Continentality is defined by low precipitation and strong seasonality of temperature. Figs. 2 and 3 reflect the history of continental
climate in central Europe by showing the patterns of MART and MAP,
both as palaeo-data alone and as differences of palaeo-minus-modern
data. Fig. 4 provides patterns of WMP that indicate the course of the
development of summer rain.
250
300
225
275
200
250
175
225
HMP [mm]
MARP [mm]
To facilitate the study of Miocene precipitation patterns in Europe,
mean annual precipitation (MAP), monthly precipitation of the driest
month (LMP), monthly precipitation of the wettest month (HMP),
monthly precipitation of the warmest month (WMP), and the mean
annual range of precipitation (MARP — the difference between HMP
and LMP) were estimated via CA. In the first step, the entire data set
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
that reflect the relationships between two parameters. Where the
correlation coefficient expresses how reliably the data are related to
one another, the regressional gradients show the nature of those
relationships.
The strongest correlation with the highest correlation coefficients
(N0.9, Table 1) in the entire data set arises between HMP and WMP.
Both parameters correlate with the other precipitation parameters, to
various degrees but always in a similar way. When the different time
intervals are compared, the correlation coefficients between HMP and
WMP are seen to increase continuously from the Middle to the Late
Miocene. Thus WMP, as estimated using CA, is not only a good measure for summer precipitation but can be assumed to reflect general
trends and patterns in precipitation parameters, especially for postLanghian times.
The graphical regressions shown in Fig. 1 indicate that the gradient
of regression between WMP and the other precipitation parameters is
steepest in Burdigalian, Serravallian and Messinian data sets. The low
number of data available for the Burdigalian stage does not allow for
further interpretation, but the generally increasing gradient in postLanghian data evidences an increase in seasonal differentiation of
precipitation towards a more pronounced precipitation peak in
summer. On the other hand, despite high absolute values for summer
precipitation and a high correlation coefficient, Langhian data show
the lowest regressional gradient, reflecting generally humid conditions with no specific wetter or drier season in summer.
150
125
200
175
100
150
75
125
100
50
50
100
50
150
1,600
60
1,400
50
1,200
40
LMP [mm]
MAP [mm]
100
150
WMP [mm]
WMP [mm]
1,000
30
20
800
10
600
0
50
100
WMP [mm]
150
50
100
150
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
Table 1
Correlation coefficients between all precipitation parameters (Pearsons r) for each
Miocene stage; most significant values N 0.9 are given in bold.
MAP
Messinian (n = 38)
HMP
0.403021
LMP
0.531935
WMP
0.303088
MARP
0.212944
MART
− 0.140436
HMP
LMP
WMP
0.696901
0.982339
− 0.521682
− 0.440914
0.650542
− 0.208308
− 0.153133
− 0.672101
− 0.479250
MARP
3.3. Seasonality gradients
0.740779
0.979634
− 0.478946
− 0.672349
0.736951
− 0.404016
− 0.471745
− 0.645457
− 0.742259
0.687520
Serravallian (n = 40)
HMP
0.551581
LMP
0.571816
WMP
0.513372
MARP
− 0.031048
MART
− 0.336340
0.641429
0.973449
− 0.237586
− 0.493868
0.594771
− 0.026666
− 0.481734
− 0.453626
− 0.465477
0.052502
Langhian (n = 41)
HMP
0.305209
LMP
0.361196
WMP
0.315189
MARP
− 0.175075
MART
− 0.026423
0.441367
0.952162
− 0.324336
− 0.155001
0.390299
− 0.050431
0.130017
− 0.597894
− 0.238667
0.332205
Burdigalian (n = 14)
HMP
0.557893
LMP
0.381232
WMP
0.536028
MARP
− 0.308981
MART
− 0.533929
0.699585
0.970741
− 0.587317
− 0.793751
0.664921
− 0.363037
− 0.712488
− 0.764483
− 0.735265
0.347334
All data
HMP
LMP
WMP
MARP
MART
− 0.465211
0.605831
0.972750
− 0.509534
− 0.177360
− 0.657772
0.376803
0.561957
− 0.379528
The differences between modern and palaeo-data document the
very low degree of change in precipitation (MAP, WMP) and MART
during the Miocene compared to the transformations that occurred
post-Miocene and up to the present. Although absolute values are
clearly changing with the approach of the Messinian, they are still
remote from the modern situation.
0.440318
Tortonian (n = 36)
HMP
0.608278
LMP
0.315546
WMP
0.520197
MARP
0.039588
MART
− 0.279421
0.408527
0.041926
0.483416
0.336634
− 0.248167
205
To aid in analysis of the latitudinal gradients of precipitation and
MART as measures of continentality, fossil climate data are plotted
against longitude in Fig. 5, while correlation coefficients are given in
Table 2. Here none of the data reveal statistically significant
longitudinal gradients. Only Messinian and Burdigalian data (with a
small data sample) show a correlation between longitude and WMP,
and Messinian and Serravallian data to some extent with MAP.
The lack of significant correlation between longitude and MART
can mainly be traced to the aforementioned buffering effect of the
Paratethys, which suppressed any development towards temperature
seasonality in the Pannonian Basin. Nevertheless, data from the
Messinian stage show a slight, although not statistically significant,
increase in MART, together with the strongest decrease in MAP and
WMP from west to east of all data. With its combination of
characteristics, the Messinian stands alone as the only stage to display
a tendency towards higher continentality in Eastern Europe. All other
Miocene stages show either no longitudinal changes, or not in this
combination. In short, evidence for continental climate in Eastern
Europe first appears with the Messinian.
4. Discussion
4.1. Miocene climate
− 0.532256
The mean annual range of temperature (MART, Fig. 2) is lowest
during Burdigalian and Langhian in Western Europe and increases
beginning with the Middle Miocene. However, the coastal parts of the
continent display only minor changes, reflecting a persistent low
seasonality of temperature influenced by the Atlantic Ocean. This
pattern is the first evidence for a Neogene climatic differentiation
between oceanic and continental climate in Europe. Due to the buffering effect of the large Paratethys sea, temperatures in Eastern
Europe also remain equable at least until the Tortonian (Bruch et al.,
2004, 2006, 2007). During the Miocene, MART patterns were evidently not as strictly east–west oriented as they are today.
Miocene precipitation data all show generally humid conditions
for the entire Miocene. Besides a general decrease in mean annual
precipitation (MAP, Fig. 3), the Langhian and Tortonian appear to be
wetter phases compared to the times before and after. Moreover, a
progressive spatial differentiation appears in the late Middle Miocene
(Serravallian) with lower precipitation in the eastern part of central
Europe.
Summer precipitation (WMP, Fig. 4) decreases beginning with the
Middle Miocene, and all stages to some extent show a longitudinal
differentiation, with slightly lower values in eastern Europe. This
trend becomes more obvious towards the Late Miocene and is the
most prominent observed pattern change of all precipitation parameters studied. Beyond changes in temperature, shifts in the annual
distribution of precipitation may have played a major role in postLanghian climate changes.
To date, interpretation of Miocene climate has mainly been based
on temperature parameters, with Miocene rainfall generally described
in terms of mean annual precipitation. The current study relies on a
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
present values. General climatic development over time can be summarised as follows.
Langhian data show very high humidity, with a peak in precipitation that does not match summer precipitation and therefore
occurred other than in summer, and no longitudinal gradient.
Serravallian data show slightly less humid conditions, evidence of
a summer peak in precipitation, and no significant longitudinal
gradient.
Tortonian data evidence slightly higher humidity than in previous
times with increasing evidence of a summer peak in precipitation, but
no longitudinal gradients.
Messinian data evidence less humid conditions, increasing
evidence of a summer peak in precipitation, and first evidence of
longitudinal gradients in WMP and MAP.
This succession testifies to a general decrease in precipitation since
the Middle Miocene and the onset of continental climate conditions in
Eastern Europe in the Late Miocene.
Beyond the general decrease in MAP, the Langhian and Tortonian
appear to have been wetter than the stages before and after. Wetter
conditions during the Langhian may be related to the Mid Miocene
Climatic Optimum (MMCO) and are widely discussed in the literature
(e.g., Kürschner et al., 2008; Retallack, 2009; Wan et al., 2009; You
et al., 2009). It is worth noting for general discussion of MMCO that
our data show a strong annual range of precipitation in central Europe
together with the highest values for all precipitation parameters,
reflecting generally very humid conditions with some seasonality, but
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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.,
2007).
The prevalence of wetter conditions during the Tortonian than in
preceding and successive stages has been less thoroughly discussed,
mainly because the focus so far has been on comparing Middle and
Late Miocene conditions and on the general decrease in humidity.
Only Böhme et al. (2008, and 2011-this issue) describe wetter
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
with our results. On the other hand, their data indicate lower than
present MAP during short-term dryer phases, where our data do not
confirm such low values for any of the parameters analysed. This may
reflect the challenge of reconstructing very dry conditions on the basis
of plant fossils (see Böhme et al., 2006, 2007). The lack of fossil floras
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preserved under dry conditions causes a strong bias in our data set with
gaps towards southern Europe, where plant proxy data are available
only from moister regions of Spain (NW coast) and Italy (N Italy).
Moreover, in central and eastern Europe as well, faunas and floras
usually come from different stratigraphic levels and taphonomic
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), is confirmed by our data. That signal is
revealed in our data with especial clarity in the increasing longitudinal
gradients of MAP and WMP over time (Fig. 5).
Miocene climatic changes after MMCO are testified in our data
set by three major factors: (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.
4.2. Landscape opening in Europe
The expansion of open landscapes during the Miocene in Europe
has been the subject of widespread and sometimes heated debate
based on fossil fauna and flora. Strömberg et al. (2007) provide a
comprehensive review of the discussion. Large mammal data speak
for the presence of open environments in southern Europe since the
early Late Miocene (e.g., Agustí et al., 1999; Fortelius et al., 2006).
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
fossils, only some data on phytoliths and pollen favour grassdominated savannas or open woodlands. Showing that such habitats
were widely established in the eastern parts of southern Europe
during the Late Miocene based on phytolith analyses, Strömberg et al.
(2007) propose that relatively open habitats developed in Asia Minor
beginning already in the Early Miocene. For Akgün et al. (2007),
however, analysis of Anatolian pollen profiles documents increasing
abundance of open vegetation taxa only during the Tortonian. In
Fig. 5. Longitudinal gradients of MART, MAP, and WMP; red circle — Burdigalian, blue
triangle — Langhian; green reversed triangle — Serravallian; purple square — Tortonian;
yellow rhombus — Messinian. Correlation coefficients in Table 2.
addition, Syabryaj et al. (2007) describe the vegetation development
of the Ukrainian plain as a stepwise opening of the forests, with the
first steppe grasslands arising during the early Tortonian (Khersonian) and expanding considerably during late Messinian (Pontian).
The same pattern of development with an onset of opening of
vegetation at the Khersonian stage is observed by Shatilova et al.
(2010) for a shallow-marine succession in eastern Georgia. Only
Kovar-Eder et al. (2008) provide a broader view of the European
vegetation history, based on a quantitative analysis of a huge floristic
data set from Europe. The authors summarise that for the Tortonian,
Table 2
Correlation coefficients between longitude and climate parameters (Pearsons r) for
each Miocene stage.
Messinian (n = 38)
Tortonian (n = 36)
Serravallian (n = 41)
Langhian (n = 40)
Burdigalian (n = 14)
MART
MAP
WMP
0.2595
− 0.0862
0.1782
0.2717
0.5436
− 0.3517
0.2434
− 0.4078
− 0.1932
0.0551
− 0.492
0.1649
− 0.1312
− 0.0154
− 0.4388
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“first records of xeric grasslands are found along the northern margin
of the Black Sea”, whereas during the Messinian “open woodlands
increasingly appeared in central and southern parts of Italy and
Greece”.
Those patterns are in agreement with our observation of the
earliest data to support continental climate in Eastern Europe during
the Messinian. Our data cannot be correlated with an onset of opening
of vegetation during or prior to the Tortonian. Either that development in vegetation is not related to the climate parameters analysed
here, or, conceivably other maybe non climatic parameters played a
role, as discussed controversially by various authors for example for
atmospheric CO2 concentration (e.g., Pagani et al., 1999; Cowling,
1999; Kürschner et al., 2008) or for the coevolution of grasses and
grazers (Retallack, 2001). It is also possible that the implications of
certain taxa as evidence for dryness and openness (e.g., Poaceae and
Asteraceae) are overestimated by qualitative approaches or underestimated by our method due to the usually wide climatic ranges of
these taxa. Hence, our data may lack the required resolution and/or
spatial coverage to fully decipher the influence of continentality on
vegetation and to correlate climatic and vegetation data statistically.
This could apply especially to southern and Eastern Europe with their
still scanty data coverage, but also points back to the methodological
question of the ability of plant proxies to detect dry events, as
discussed above.
Nevertheless, climatic data that can quantify continentality are a
valuable basis for explaining the expansion of grassland in Eurasia.
Further studies on a broader geographic scale, with an enhanced focus
on southern Europe and central Eurasia, will aid in quantifying the
development of precipitation patterns and understanding their
influence on the history of landscape opening during the Neogene.
Supplementary materials related to this article can be found,
online, at doi: 10.1016/j.palaeo.2010.10.002.
Acknowledgements
We would like to express our gratitude to all our colleagues in the
NELCIME network who contributed their data, expertise and discussion. We very much appreciate the opportunity we have had to work
with them over the last ten years and our many stimulating and
fruitful discussions with various NECLIME members during annual
meetings and on other occasions. We are also thankful to the two
anonymous reviewers who helped to considerably improve the
quality of the manuscript. Special thanks go to Dr. Christopher Traiser
(Tübingen), who invested much effort in making the data available on
Pangaea.
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