The origin of grasslands in the temperate forest zone of east-central Europe: long-term legacy of climate and human impact.

The post-glacial fate of central European grasslands has stimulated palaeoecological debates for a century. Some argued for the continuous survival of open land, while others claimed that closed forest had developed during the Middle Holocene. The reasons behind stability or changes in the proportion of open land are also unclear. We aim to reconstruct regional vegetation openness and test the effects of climate and human impact on vegetation change throughout the Holocene. We present a newly dated pollen record from north-western fringes of the Pannonian Plain, east-central Europe, and reconstruct Holocene regional vegetation development by the REVEALS model for 27 pollen-equivalent taxa. Estimated vegetation is correlated in the same area with a human activity model based on all available archaeological information and a macrophysical climate model. The palaeovegetation record indicates the continuous presence of open land throughout the Holocene. Grasslands and open woodlands were probably maintained by local arid climatic conditions during the early Holocene delaying the spread of deciduous (oak) forests. Significantly detectable human-made landscape transformation started only after 2000 BC. Our analyses suggest that Neolithic people spread into a landscape that was already open. Humans probably contributed to the spread of oak, and influenced the dynamics of hazel and hornbeam.

[1]  B. Huntley,et al.  Holocene persistence of wooded steppe in the Great Hungarian Plain , 2010 .

[2]  Paul D. Henne,et al.  Did soil development limit spruce (Picea abies) expansion in the Central Alps during the Holocene? Testing a palaeobotanical hypothesis with a dynamic landscape model , 2011 .

[3]  Jan Kolář,et al.  Spatio-temporal modelling as a way to reconstruct patterns of past human activities. , 2016, Archaeometry.

[4]  J. Svenning,et al.  A review of natural vegetation openness in north-western Europe , 2002 .

[5]  K. Behre Evidence for Mesolithic agriculture in and around central Europe? , 2006 .

[6]  Spassimir Tonkov,et al.  Climate variability and associated vegetation response throughout Central and Eastern Europe (CEE) between 60 and 8 ka , 2014 .

[7]  M. Horsák,et al.  Holocene history of a Cladium mariscus-dominated calcareous fen in Slovakia: vegetation stability and landscape development , 2013 .

[8]  A. Mercuri,et al.  Mediterranean and north-African cultural adaptations to mid-Holocene environmental and climatic changes , 2011 .

[9]  K. Hjelle,et al.  Ecology and long-term land-use, palaeoecology and archaeology – the usefulness of interdisciplinary studies for knowledge-based conservation and management of cultural landscapes , 2012 .

[10]  H. Birks,et al.  Pollen‐based quantitative reconstructions of Holocene regional vegetation cover (plant‐functional types and land‐cover types) in Europe suitable for climate modelling , 2015, Global change biology.

[11]  J. Cumming,et al.  Where, When, and How , 2007, Research quarterly for exercise and sport.

[12]  G. Schmiedl,et al.  Massive perturbation in terrestrial ecosystems of the Eastern Mediterranean region associated with the 8.2 kyr B.P. climatic event , 2009 .

[13]  S. Sugita,et al.  The REVEALS model, a new tool to estimate past regional plant abundance from pollen data in large lakes: validation in southern Sweden , 2008 .

[14]  V. Ložek,et al.  Mollusc succession of a prehistoric settlement area during the Holocene: A case study of the České středohoří Mountains (Czech Republic) , 2013 .

[15]  C. Leff Czech and Slovak Republics , 1996 .

[16]  V. Pavlů,et al.  Origin and history of grasslands in Central Europe – a review , 2013 .

[17]  E. Eckmeier,et al.  Pedogenesis of Chernozems in Central Europe — A review , 2007 .

[18]  D. Zelený,et al.  High species richness in hemiboreal forests of the northern Russian Altai, southern Siberia , 2012 .

[19]  H. Birks,et al.  Holocene changes in vegetation composition in northern Europe: why quantitative pollen-based vegetation reconstructions matter , 2014 .

[20]  M. Gaillard Pollen Methods and Studies : Archaeological applications , 2007 .

[21]  V. Ložek Molluscan fauna from the loess series of Bohemia and Moravia , 2001 .

[22]  Thomas Raddatz,et al.  A reconstruction of global agricultural areas and land cover for the last millennium , 2008 .

[23]  H. Birks,et al.  Spatial structure of the 8200 cal yr BP event in northern Europe , 2007 .

[24]  A. Lotter,et al.  Central European vegetation response to abrupt climate change at 8.2 ka , 2001 .

[25]  Shawn A. Ross,et al.  Environmental conditions in the SE Balkans since the Last Glacial Maximum and their influence on the spread of agriculture into Europe , 2013 .

[26]  Rob Marchant,et al.  Neolithic agriculture on the European western frontier: the boom and bust of early farming in Ireland , 2014 .

[27]  J. Christen,et al.  Flexible paleoclimate age-depth models using an autoregressive gamma process , 2011 .

[28]  A. Molnár,et al.  The late quaternary paleoecology and environmental history of Hortobágy, a unique mosaic alkaline steppe from the heart of the Carpathian Basin , 2013 .

[29]  B. MacFadden,et al.  Seasonal and geographic climate variabilities during the Last Glacial Maximum in North America: Applying isotopic analysis and macrophysical climate models , 2009 .

[30]  M. Horsák,et al.  Early occurrence of temperate oak-dominated forest in the northern part of the Little Hungarian Plain, SW Slovakia , 2014 .

[31]  F. Vera Grazing Ecology and Forest History , 2000 .

[32]  P. Kuneš,et al.  Soil phosphorus as a control of productivity and openness in temperate interglacial forest ecosystems , 2011 .

[33]  Philippe Crombé,et al.  14C dates as demographic proxies in Neolithisation models of northwestern Europe: a critical assessment using Belgium and northeast France as a case-study , 2014 .

[34]  Andrew S. Fairbairn,et al.  Neolithic human impact on the landscapes of North-East Hungary inferred from pollen and settlement records , 2012, Vegetation History and Archaeobotany.

[35]  H. Ellenberg Vegetation Mitteleuropas mit den Alpen : in ökologischer ,dynamischer und historischer Sicht , 2010 .

[36]  P. Sümegi,et al.  Woodland-Grassland ecotonal shifts in environmental mosaics: Lessons learnt from the environmental history of the carpathian basin (Central Europe) during the holocene and the last ice age based on investigation of paleobotanical and mollusk remains , 2012 .

[37]  Thomas Raus,et al.  Karte der natürlichen Vegetation Europas / Map of the Natural Vegetation of Europe - Maßstab / Scale 1:2,500,000 , 2000 .

[38]  Han Olff,et al.  Shifting mosaics in grazed woodlands driven by the alternation of plant facilitation and competition , 1999 .

[39]  R. Kozáková,et al.  Relative pollen productivity estimates in the modern agricultural landscape of Central Bohemia (Czech Republic) , 2012 .

[40]  K. Behre,et al.  interpretation of anthropogenic indicators in pollen diagrams , 1981 .

[41]  L. Kalniņa,et al.  Annual pollen traps reveal the complexity of climatic control on pollen productivity in Europe and the Caucasus , 2010 .

[42]  S. Sugita,et al.  Theory of quantitative reconstruction of vegetation I: pollen from large sites REVEALS regional vegetation composition , 2007 .

[43]  A. Lister,et al.  A long-term perspective on ungulate–vegetation interactions , 2003 .

[44]  R Core Team,et al.  R: A language and environment for statistical computing. , 2014 .

[45]  P. Pokorný,et al.  Detection of the impact of early Holocene hunter-gatherers on vegetation in the Czech Republic, using multivariate analysis of pollen data , 2008 .

[46]  Marc D. Abrams,et al.  Fire and the Development of Oak Forests , 1992 .

[47]  P. Pokorný,et al.  Palaeoenvironmental research of the Schwarzenberg Lake, southern Bohemia, and exploratory excavations of this key Mesolithic archaeological area Paleoenvironmentální výzkum zaniklého jezera Schwarzenberg v jižních Čechách a navazující průzkum mezolitického archeologického prostoru , 2010 .

[48]  John Meadows,et al.  Summed radiocarbon calibrations as a population proxy: a critical evaluation using a realistic simulation approach , 2014 .

[49]  K. Bennett,et al.  The Neolithic transition - fact or fiction? Palaeoecological evidence from the Balkans , 1994 .

[50]  D. Schwartz,et al.  Vegetation history of chernozems in the Czech Republic , 2014, Vegetation History and Archaeobotany.

[51]  A. Mercuri,et al.  ANTHROPOGENIC POLLEN INDICATORS (API) FROM ARCHAEOLOGICAL SITES AS LOCAL EVIDENCE OF HUMAN-INDUCED ENVIRONMENTS IN THE ITALIAN PENINSULA , 2013 .

[52]  F. Firbas Spät- und nacheiszeitliche Waldgeschichte Mitteleuropas nördlich der Alpen , 1950 .

[53]  P. Szabó,et al.  Continuity and change in the vegetation of a Central European oakwood , 2013 .

[54]  Kamil Rybníček,et al.  Erste Ergebnisse paläogeobotanischer Untersuchungen des Moores bei Vracov, Südmähren , 1972, Folia Geobotanica et Phytotaxonomica.

[55]  J. L. Reveal,et al.  Ancient Woodland, Its History, Vegetation and Uses in England , 1980 .

[56]  G. Matys,et al.  Šúr, a former late-glacial and Holocene lake at the westernmost margin of the Carpathians Šúr - zaniklé pozdně glaciální a holocenní jezero na nejzápadnějším okraji Karpat , 2013 .

[57]  S. Shennan,et al.  radiocarbon-inferred demographic change from Central Europe Is Neolithic land use correlated with demography ? An evaluation of pollen-derived land cover and , 2014 .

[58]  S. Riehl,et al.  Local Holocene environmental indicators in Upper Mesopotamia: Pedogenic carbonate record vs. archaeobotanical data and archaeoclimatological models , 2009 .

[59]  A. Havinga A palynological investigation in the pannonian climate region of lower Austria , 1972 .

[60]  G. Wightman The Czech and Slovak Republics , 1993 .

[61]  Martin Hinz,et al.  Demography and the intensity of cultural activities: an evaluation of Funnel Beaker Societies (4200–2800 cal BC) , 2012 .

[62]  H. Birks Mind the gap: how open were European primeval forests? , 2005, Trends in ecology & evolution.

[63]  N. Benecke,et al.  Holocene survival of the wild horse in Europe: a matter of open landscape? , 2011 .

[64]  R. Betts,et al.  Land use/land cover changes and climate: modeling analysis and observational evidence , 2011 .

[65]  K. Bennett,et al.  Determination of the number of zones in a biostratigraphical sequence. , 1996, The New phytologist.

[66]  Tim Kerig,et al.  Regional population collapse followed initial agriculture booms in mid-Holocene Europe , 2013, Nature Communications.

[67]  F. Mitchell How open were European primeval forests? Hypothesis testing using palaeoecological data , 2005 .

[68]  Petr Kočár,et al.  Trends in cereal cultivation in the Czech Republic from the Neolithic to the Migration period (5500 b.c.–a.d. 580) , 2013, Vegetation History and Archaeobotany.

[69]  F. Mazier,et al.  Testing the effect of site selection and parameter setting on REVEALS-model estimates of plant abundance using the Czech Quaternary Palynological Database , 2012 .

[70]  B. Shuman Patterns, processes, and impacts of abrupt climate change in a warm world: the past 11,700 years , 2012 .

[71]  M. Horsák,et al.  Modern analogues from the Southern Urals provide insights into biodiversity change in the early Holocene forests of Central Europe , 2010 .

[72]  T. Reitalu,et al.  The role of landscape structure in determining palynological and floristic richness , 2012, Vegetation History and Archaeobotany.

[73]  Thomas Giesecke,et al.  Holocene land-cover reconstructions for studies on land cover-climate feedbacks , 2010 .

[74]  M. Chytrý,et al.  Mid-Holocene bottleneck for central European dry grasslands: Did steppe survive the forest optimum in northern Bohemia, Czech Republic? , 2015 .

[75]  O. Kovárík,et al.  : Czech Quaternary Palynological Database - PALYCZ: review andbasic statistics of the data , 2009 .

[76]  M. Regnell Plant subsistence and environment at the Mesolithic site Tågerup, southern Sweden: new insights on the “Nut Age” , 2011, Vegetation History and Archaeobotany.

[77]  O. Moine,et al.  High-resolution record of the environmental response to climatic variations during the Last Interglacial-Glacial cycle in Central Europe: the loess-palaeosol sequence of Dolní Věstonice (Czech Republic) , 2013 .

[78]  A. Wiersma,et al.  Model–data comparison for the 8.2 ka BP event: Confirmation of a forcing mechanism by catastrophic drainage of Laurentide Lakes. , 2006 .

[79]  A. Lotter,et al.  Mesolithic agriculture in Switzerland? A critical review of the evidence , 2007 .

[80]  K. Willis,et al.  DOES SOIL CHANGE CAUSE VEGETATION CHANGE OR VICE VERSA? A TEMPORAL PERSPECTIVE FROM HUNGARY , 1997 .

[81]  Thomas Giesecke,et al.  Quantitative reconstructions of changes in regional openness in north-central Europe reveal new insights into old questions , 2012 .

[82]  A. Mercuri Genesis and evolution of the cultural landscape in central Mediterranean: the ‘where, when and how’ through the palynological approach , 2014, Landscape Ecology.

[83]  H. Ellenberg,et al.  Vegetation Mitteleuropas mit den Alpen , 1984 .

[84]  Benjamin Smith,et al.  Regional climate model simulations for Europe at 6 and 0.2 k BP: sensitivity to changes in anthropogenic deforestation , 2013 .

[85]  H. Birks,et al.  The pace of Holocene vegetation change : testing for synchronous developments , 2011 .

[86]  T. Hickler,et al.  Origin of the forest steppe and exceptional grassland diversity in Transylvania (central‐eastern Europe) , 2015 .

[87]  J. Kaplan,et al.  The prehistoric and preindustrial deforestation of Europe , 2009 .

[88]  Christopher Bronk Ramsey,et al.  BAYESIAN ANALYSIS OF RADIOCARBON DATES , 2009 .

[89]  C. Buck,et al.  IntCal13 and Marine13 Radiocarbon Age Calibration Curves 0–50,000 Years cal BP , 2013, Radiocarbon.

[90]  B. Geel Non-Pollen Palynomorphs , 2002 .

[91]  S. Sugita A Model of Pollen Source Area for an Entire Lake Surface , 1993, Quaternary Research.

[92]  Dagmar Dreslerová,et al.  Human Response to Potential Robust Climate Change around 5500 cal BP in the Territory of Bohemia (the Czech Republic) , 2012 .

[93]  K. Willis,et al.  Do dung fungal spores make a good proxy for past distribution of large herbivores , 2013 .

[94]  Sarah L. R. Mason Fire and Mesolithic subsistence — managing oaks for acorns in northwest Europe? , 2000 .

[95]  P. Kuneš,et al.  Present-Day Vegetation Helps Quantifying Past Land Cover in Selected Regions of the Czech Republic , 2014, PloS one.

[96]  H. Joosten,et al.  Corylus expansion and persistent openness in the early Holocene vegetation of northern central Europe , 2014 .

[97]  H. Weiss,et al.  Formal Subdivision of the Holocene Series/Epoch , 2014 .

[98]  T. Kull,et al.  Palynological richness and pollen sample evenness in relation to local floristic diversity in southern Estonia , 2011 .