Beringia as a glacial refugium for boreal trees and shrubs: new perspectives from mapped pollen data

Aim Beringia, far north‐eastern Siberia and north‐western North America, was largely unglaciated during the Pleistocene. Although this region has long been considered an ice‐age refugium for arctic herbs and shrubs, little is known about its role as a refugium for boreal trees and shrubs during the last glacial maximum (LGM, c. 28,000–15,000 calibrated years before present). We examine mapped patterns of pollen percentages to infer whether six boreal tree and shrub taxa (Populus, Larix, Picea, Pinus, Betula, Alnus/Duschekia) survived the harsh glacial conditions within Beringia. Methods Extensive networks of pollen records have the potential to reveal distinctive temporal–spatial patterns that discriminate between local‐ and long‐distance sources of pollen. We assembled pollen records for 149 lake, peat and alluvial sites from the Palaeoenvironmental Arctic Sciences database, plotting pollen percentages at 1000‐year time intervals from 21,000 to 6000 calibrated years before present. Pollen percentages are interpreted with an understanding of modern pollen representation and potential sources of long‐distance pollen during the glacial maximum. Inferences from pollen data are supplemented by published radiocarbon dates of identified macrofossils, where available. Results Pollen maps for individual taxa show unique temporal‐spatial patterns, but the data for each taxon argue more strongly for survival within Beringia than for immigration from outside regions. The first increase of Populus pollen percentages in the western Brooks Ranges is evidence that Populus trees survived the LGM in central Beringia. Both pollen and macrofossil evidence support Larix survival in western Beringia (WB), but data for Larix in eastern Beringia (EB) are unclear. Given the similar distances of WB and EB to glacial‐age boreal forests in temperate latitudes of Asia and North America, the widespread presence of Picea pollen in EB and Pinus pollen in WB indicates that Picea and Pinus survived within these respective regions. Betula pollen is broadly distributed but highly variable in glacial‐maximum samples, suggesting that Betula trees or shrubs survived in restricted populations throughout Beringia. Alnus/Duschekia percentages show complex patterns, but generally support a glacial refugium in WB. Main conclusions Our interpretations have several implications, including: (1) the rapid post‐glacial migration rate reported for Picea in western Canada may be over estimated, (2) the expansion of trees and shrubs within Beringia should have been nearly contemporaneous with climatic change, (3) boreal trees and shrubs are capable of surviving long periods in relatively small populations (at the lower limit of detection in pollen data) and (4) long‐distance migration may not have been the predominant mode of vegetation response to climatic change in Beringia.     

The mid‐Holocene vegetation of the Mediterranean region and southern Europe,and comparison with the present day

Aim To contribute to the intense debate surrounding the relative influence of climate and humans on Mediterranean‐region land cover over the past 6000 years, we assess the Holocene biogeography and vegetation history of southern Europe by means of an extensive pollen record dataset. Location The Mediterranean biogeographical zone and neighbouring parts of Iberia, the Alps and Anatolia, between 30° N, 48° N, 10° W and 45° E. Methods We compiled a southern European pollen record dataset using available pollen databases (124 sites) and other sources (74 sites), with improved spatial coverage and dating control compared with earlier studies. We used only those sites that had pollen data for both 0 ka and 6 ka. We reconstructed mid‐Holocene and present‐day biomes, arboreal pollen percentages and distribution and relative abundance of 11 key woody taxa, with anomaly maps. Results Northern temperate forest biomes extended further south at the mid‐Holocene than at present, but not as far as earlier studies suggested. Sclerophyllous vegetation occurred along the Mediterranean coast throughout the region at 6 ka. Arboreal pollen percentages were up to 50% higher than at present. At 6 ka, Olea, Fagus and Juniperus had smaller distributions and/or abundances; Abies, Cedrus and both deciduous and evergreen Quercus had larger distributions and/or abundances; Phillyrea, Pistacia and Cistus showed minimal difference; and Pinus showed a cosmopolitan distribution with variable abundance. Main conclusions Temporal difference analysis is more meaningful when only sites containing samples for all time slices are analysed. During the mid‐Holocene, southern Europe was more heavily forested with temperate vegetation than it is at present, but drought‐tolerant xeric vegetation was still widespread along the southern margins of the region. Although human land use may have caused the degradation of land between the mid‐Holocene and the present, the mere presence of xeric vegetation in the Mediterranean region does not require human impact. This challenges the commonly held belief that modern Mediterranean vegetation represents a ‘degraded’ state.     

Tree line identification from pollen data: beyond the limit?

Aim The boreal tree line is a prominent biogeographic feature, the position of which reflects climatic conditions. Pollen is the key sensor used to reconstruct past tree line patterns. Our aims in this study were to investigate pollen–vegetation relationships at the boreal tree line and to assess the success of a modified version of the biomization method that incorporates pollen productivity and dispersal in distinguishing the tree line. Location Northern Canada (307 sites) and Alaska (316 sites). Methods The REVEALS method for estimating regional vegetation composition from pollen data was simplified to provide correction factors to account for differential production and dispersal of pollen among taxa. The REVEALS‐based correction factors were used to adapt the biomization method and applied as a set of experiments to pollen data from lake sediments and moss polsters from the boreal tree line. Proportions of forest and tundra predicted from modern pollen samples along two longitudinal transects were compared with those derived from a vegetation map by: (1) a tally of ‘correct’ versus ‘incorrect’ assignments using vegetation in the relevant map pixels, and (2) a comparison of the shape and position of north–south forest‐cover curves generated from all transect pixels and from pollen data. Possible causes of bias in the misclassifications were assessed. Results Correcting for pollen productivity alone gave fewest misclassifications and the closest estimate of the modern mapped tree line position (Canada, + 300 km; Alaska, + 10 km). In Canada success rates were c. 40–70% and all experiments over‐predicted forest cover. Most corrections improved results over uncorrected biomization; using only lakes improved success rates to c. 80%. In Alaska success rates were 70–80% and classification errors were more evenly distributed; there was little improvement over uncorrected biomization. Main conclusions Corrected biomization should improve broad‐scale reconstructions of spatial patterns in forest/non‐forest vegetation mosaics and across climate‐sensitive ecotones. The Canadian example shows this is particularly the case in regions affected by taxa with extremely high pollen productivity (such as Pinus). Improved representation of actual vegetation distribution is most likely if pollen data from lake sediments are used because the REVEALS algorithm is based on the pollen dynamics of lake‐based systems.     

Effects of the sampling design and selection of parameter values on pollen-based quantitative reconstructions of regional vegetation: a case study in southern Sweden using the REVEALS model

The need for quantification of land cover from pollen data has led to the development of a Landscape Reconstruction Algorithm (LRA). The LRA includes several models of which the REVEALS model estimates regional vegetation abundance using pollen assemblages from large sites (lakes or bogs). In this paper we explore the effects of selection and number of pollen samples, and choice of pollen productivity estimates on the REVEALS results. The effect of the size of vegetation surveys is also tested. The results suggest that the differences between two sizes of vegetation surveys have little effect on the model validation. The “characteristic radius” of regional vegetation in southern Sweden was estimated as 200 km. However, the vegetation composition in a 100 × 100 km² square matches well with that estimated by REVEALS. Whether 25, 20 (outliers excluded) or 4 pollen samples are used does not change the REVEALS reconstructions much although the error estimates are larger when outliers are included, and very large when only four samples are used. Therefore validation of the REVEALS model and REVEALS reconstructions of past vegetation can be performed using a limited number of pollen samples, although with caution. The use of many pollen samples from multiple sites is always better whenever possible. REVEALS reconstructions are closer to the actual vegetation when the Danish Pollen Productivity Estimates (PPEs) are used instead of the Swedish PPEs for Cereals, Rumex acetosa/acetosella, Plantago lanceolata and Calluna, indicating that the Danish PPEs are more reliable than the Swedish ones for those taxa. It is recommended to test more than one set of PPEs in validation and applications of the REVEALS model for a better evaluation of the results.     

Are pollen records from small sites appropriate for REVEALS model-based quantitative reconstructions of past regional vegetation? An empirical test in southern Sweden

In this paper we test the performance of the Regional Estimates of VEgetation Abundance from Large Sites (REVEALS) model using pollen records from multiple small sites. We use Holocene pollen records from large and small sites in southern Sweden to identify what is/are the most significant variable(s) affecting the REVEALS-based reconstructions, i.e. type of site (lakes and/or bogs), number of sites, site size, site location in relation to vegetation zones, and/or distance between small sites and large sites. To achieve this objective we grouped the small sites according to (i) the two major modern vegetation zones of the study region, and (ii) the distance between the small sites and large lakes, i.e. small sites within 50, 100, 150, or 200 km of the large lakes. The REVEALS-based reconstructions were performed using 24 pollen taxa. Redundancy analysis was performed on the results from all REVEALS-model runs using the groups within (i) and (ii) separately, and on the results from all runs using the groups within (ii) together. The explanatory power and significance of the variables were identified using forward selection and Monte Carlo permutation tests. The results show that (a) although the REVEALS model was designed for pollen data from large lakes, it also performs well with pollen data from multiple small sites in reconstructing the percentage cover of groups of plant taxa (e.g. open land taxa, summer-green trees, evergreen trees) or individual plant taxa; however, in the case of this study area, the reconstruction of the percentage cover of Calluna vulgaris, Cyperaceae, and Betula may be problematic when using small bogs; (b) standard errors of multiple small-site REVEALS estimates will generally be larger than those obtained using pollen records from large lakes, and they will decrease with increasing size of pollen counts and increasing number of small sites; (c) small lakes are better to use than small bogs if the total number of small sites is low; and (d) the size of small sites and the distance between them do not play a major role, but the distance between the small sites and landscape/vegetation boundaries is a determinant factor for the accuracy of the vegetation reconstructions.     

A survey of modern pollen and vegetation along a south–north transect in Mongolia

Aim This modern pollen‐rain study documents the spatial and quantitative relationships between modern pollen and vegetation in Mongolia, and explores the potential for using this relationship in palaeoclimatic reconstructions. Location East‐central Mongolia. Methods We collected 104 pollen surface samples along a south–north transect across five vegetation zones in Mongolia. Discriminant analysis was used to classify the modern pollen spectra into five pollen assemblages corresponding to the five vegetation zones. Hierarchical cluster analysis was used to divide the main pollen taxa into two major groups and seven subgroups representing the dry and moist vegetation types and the main vegetation communities within them. Results Each vegetation zone along the transect can be characterized by a distinctive modern pollen assemblage as follows: (1) desert zone: Chenopodiaceae–Zygophyllaceae–Nitraria–Poaceae pollen assemblage; (2) desert‐steppe zone: Poaceae–Chenopodiaceae pollen assemblage; (3) steppe zone: Artemisia–Aster‐type–Poaceae–Pinus Haploxylon‐type pollen assemblage; (4) forest‐steppe zone: Pinus Haploxylon‐type–Picea–Artemisia–Betula, montane forb/shrub and pteridophyte pollen assemblage; and (5) mountain taiga zone: Pinus Haploxylon‐type–Picea–Poaceae–Cyperaceae, montane forb/shrub and Pteridophyte pollen assemblage. Main conclusions Based on the ratio between the major pollen taxon groups and subgroups, we propose two pollen–climate indices that represent the precipitation and temperature conditions in the study region. When plotted along our south–north transect, the moisture indices (M) and temperature indices (T) mimic the regional gradients of precipitation and temperature across Mongolia very closely. These pollen–climate indices can be used for palaeoclimatic reconstruction based on fossil pollen data.     

From forest to farmland: pollen‐inferred land cover change across Europe using the pseudobiomization approach

Maps of continental‐scale land cover are utilized by a range of diverse users but whilst a range of products exist that describe present and recent land cover in Europe, there are currently no datasets that describe past variations over long time‐scales. User groups with an interest in past land cover include the climate modelling community, socio‐ecological historians and earth system scientists. Europe is one of the continents with the longest histories of land conversion from forest to farmland, thus understanding land cover change in this area is globally significant. This study applies the pseudobiomization method (PBM) to 982 pollen records from across Europe, taken from the European Pollen Database (EPD) to produce a first synthesis of pan‐European land cover change for the period 9000 bp to present, in contiguous 200 year time intervals. The PBM transforms pollen proportions from each site to one of eight land cover classes (LCCs) that are directly comparable to the CORINE land cover classification. The proportion of LCCs represented in each time window provides a spatially aggregated record of land cover change for temperate and northern Europe, and for a series of case study regions (western France, the western Alps, and the Czech Republic and Slovakia). At the European scale, the impact of Neolithic food producing economies appear to be detectable from 6000 bp through reduction in broad‐leaf forests resulting from human land use activities such as forest clearance. Total forest cover at a pan‐European scale moved outside the range of previous background variability from 4000 bp onwards. From 2200 bp land cover change intensified, and the broad pattern of land cover for preindustrial Europe was established by 1000 bp . Recognizing the timing of anthropogenic land cover change in Europe will further the understanding of land cover‐climate interactions, and the origins of the modern cultural landscape.     

Upland vegetation in the north-west Iberian peninsula after the last glaciation: Forest history and deforestation dynamics

This paper presents the results of pollen analyses from organic sediments of seven cores (299 spectra) in a mountainous area of the north-west Iberian peninsula. The pollen diagrams, supported by seven¹⁴C dates, are used to construct a regional pollen sequence covering the main stages of vegetation dynamics, from the last phases of the Late-glacial until the present. During the Late-glacial Interstadial an important development of cryophilous forests (Betula andPinus) was recorded, although various mesophilous and thermophilous tree elements were also present. The Younger Dryas, palynologically clearly defined, is characterized by an important reduction in tree pollen percentages and the expansion of steppe formations (Poaceae andArtemisia). At the beginning of the Holocene, there was an expansion ofQuercus and a spread of other trees, which combined to give a vegetation cover of varied composition but dominated by mixed deciduous forests. Such forest formations prevailed in these mountains until 3000 years ago, when successive deforestation phases are recorded at various times as a result of increased farming activity. The results are compared with data from other mountainous areas in the northern Iberian peninsula and southern France.     

Past vegetation dynamics in the Yellowstone region highlight the vulnerability of mountain systems to climate change

Aim

Reconstruct the long‐term ecosystem dynamics of the region across an elevational gradient as they relate to climate and local controls. In particular, we (1) describe the dominant conifers' history; (2) assess changes in vegetation composition and distribution; and (3) note periods of abrupt change versus stability as means of better understanding vegetation responses to environmental variability.

Location

Greater Yellowstone Ecosystem (GYE; USA).

Time period

16.5 ka bp ‐present.

Major taxa studied

Juniperus, Picea, Abies, Pinus, Pseudotsuga.

Methods

The vegetation reconstruction was developed from 15 pollen records. Results were interpreted based on modern pollen–vegetation relationships estimated from a suite of regression‐based approaches.

Results

Calibrated pollen data suggest that late‐glacial vegetation, dominated by shrubs and Juniperus, lacks a modern counterpart in the area. Picea, Abies and Pinus expanded at 16 ka bp in association with postglacial warming and co‐occurred in mixed‐conifer parkland/forest after 12 ka bp . This association along with Pinus contorta forest, which was present after 9 ka bp , has persisted with little change at middle and high elevations to the present day. This stability contrasts with the dynamic history of plant communities at low elevations, where shifts between parkland, steppe and forest over the last 8,000 years were likely driven by variations in effective moisture and fire.

Main conclusions

The postglacial vegetation history of the GYE highlights the dynamic nature of mountain ecosystems and informs on their vulnerability to future climate change: (1) most of the conifers have been present in the area for >12,000 years and survived climate change by adjusting their elevational ranges; (2) some plant associations have exhibited stability over millennia as a result of nonclimatic controls; and (3) present‐day forest cover is elevationally more compressed than at any time in history, probably due to the legacy of the Medieval Climate Anomaly and the Little Ice Age.     

Creating spatially continuous maps of past land cover from point estimates: A new statistical approach applied to pollen data

Reliable estimates of past land cover are critical for assessing potential effects of anthropogenic land-cover changes on past earth surface-climate feedbacks and landscape complexity. Fossil pollen records from lakes and bogs have provided important information on past natural and human-induced vegetation cover. However, those records provide only point estimates of past land cover, and not the spatially continuous maps at regional and sub-continental scales needed for climate modelling.We propose a set of statistical models that create spatially continuous maps of past land cover by combining two data sets: 1) pollen-based point estimates of past land cover (from the REVEALS model) and 2) spatially continuous estimates of past land cover, obtained by combining simulated potential vegetation (from LPJ-GUESS) with an anthropogenic land-cover change scenario (KK10). The proposed models rely on statistical methodology for compositional data and use Gaussian Markov Random Fields to model spatial dependencies in the data.Land-cover reconstructions are presented for three time windows in Europe: 0.05, 0.2, and 6 ka years before present (BP). The models are evaluated through cross-validation, deviance information criteria and by comparing the reconstruction of the 0.05 ka time window to the present-day land-cover data compiled by the European Forest Institute (EFI). For 0.05 ka, the proposed models provide reconstructions that are closer to the EFI data than either the REVEALS- or LPJ-GUESS/KK10-based estimates; thus the statistical combination of the two estimates improves the reconstruction. The reconstruction by the proposed models for 0.2 ka is also good. For 6 ka, however, the large differences between the REVEALS- and LPJ-GUESS/KK10-based estimates reduce the reliability of the proposed models. Possible reasons for the increased differences between REVEALS and LPJ-GUESS/KK10 for older time periods and further improvement of the proposed models are discussed.     

Pollen analysis of coal-bearing Miocene sedimentary rocks from the Seyitömer Basin (Kütahya), Western Anatolia

The late Early-Middle Miocene sequences of the Seyitömer Basin (western Anatolia) were palynologically investigated. Fifty-five taxa belonging to seven gymnospermous and 48 angiospermous pollen genera were identified in the 19 productive samples. Two pollen zones were recognised based on the changing abundance of individual tree taxa. Zone 1 is characterized by predominance of Pinus and Cedrus. Zone 2 is characterized by predominance of deciduous Quercus and evergreen Quercus and a marked reduction in representation of Taxodiaceae. The differences in the pollen spectra between Zone 1 and Zone 2 may reflect the global Middle Miocene cooling. These results are largely comparable to pollen data derived from the neighbouring areas. The vegetation of the Seyitömer Basin was dominated by trees. This palynological analysis reveals the existence of a swamp-forest developed in a subtropical to warm-temperate humid climate.     

Vegetation, climate and palaeoaltitude reconstructions of the Eastern Alps during the Miocene based on pollen records from Austria, Central Europe

Aim To reconstruct the flora, vegetation, climate and palaeoaltitude during the Miocene (23.03–5.33 Ma) in Central Europe. Location Six outcrop sections located in different basins of the Central Paratethys in Austria. Methods Pollen analysis was used for the reconstruction of the vegetation and climate. The altitude of the Eastern Alps that are adjacent to the Alpine Foreland and Vienna basins has been estimated using a new quantification method based on pollen data. This method uses biogeographical and climatological criteria such as the composition of the modern vegetation belts in the European mountains and Miocene annual temperature estimates obtained from fossil pollen data. Results Pollen changes from Early to Late Miocene have been observed. The vegetation during the Burdigalian and Langhian (20.43–13.65 Ma) was dominated by thermophilous elements such as evergreen trees, typical of a present‐day evergreen rain forest at low altitudes (i.e. south‐eastern China). During the Serravallian and Tortonian (13.65–7.25 Ma) several thermophilous elements strongly decreased, and some disappeared from the Central European region. This kind of vegetation was progressively substituted by one enriched in deciduous and mesothermic plants. Middle‐altitude (Cathaya, Cedrus and Tsuga) and high‐altitude (Abies and Picea) conifers increased considerably during the Langhian and later on during the Serravallian and Tortonian. Main conclusions Pollen changes are related to climatic changes and to the uplift of the Alpine massifs. The vegetation during the Burdigalian and Langhian reflects the Miocene climatic optimum. The decrease in thermophilous plants during the Serravallian and Tortonian can be interpreted as a climatic cooling and can be correlated with global and regional climatic changes. This study shows that the palaeoaltitude of the eastern part of the Eastern Alps during the Burdigalian was not high enough for Abies and Picea to form a forest. Therefore, we inferred that the summits of most of the mountains would have been less than 1800 m. The substantial increase of middle‐ and high‐altitude conifers in the pollen spectra suggests that the uplift rate increased during the Langhian in this region. Based on higher palaeoaltitude estimations for the pollen floras from the studied sections of Austria, we infer that the uplift of the easternmost part of the Alpine chain continued during the Serravallian and Tortonian.     

Remotely‐sensed foliage cover and ground‐measured stand attributes are complimentary when estimating tree hollow abundances across relictual woodlands in agricultural landscapes

Hollows, also known as tree cavities, are critical to the survival of many animal species but are too poorly mapped across landscapes to allow for their adequate consideration in regional planning. Managing cost is important, so we tested whether freely available satellite‐derived foliage projective cover and field‐measured stand attributes could be used separately or combined to predict tree hollow abundance in relictual Australian temperate woodlands. Satellite‐derived foliage projective cover revealed variation in woody vegetation densities both within mapped woodland remnants and cleared areas of the agricultural matrix. Plot‐based field assessment of the actual number of hollows in each one‐hectare site (n = 110 sites) revealed a relationship with foliage cover. Improvement of the model was achieved if site‐based estimates of the proportion of the canopy due to Eucalyptus species and the number of mature trees per hectare were included. Remotely sensed foliage cover can improve on traditional vegetation mapping for predicting hollow‐bearing tree and hollow abundances at landscape scales when managing hollow‐dependent fauna habitat across relictual woodlands in temperate Australian agricultural landscapes. At finer scales, the addition of other predictors is necessary to raise the accuracy of the predicted hollow densities.     

A comparison of sedimentary DNA and pollen from lake sediments in recording vegetation composition at the Siberian treeline

Reliable information on past and present vegetation is important to project future changes, especially for rapidly transitioning areas such as the boreal treeline. To study past vegetation, pollen analysis is common, while current vegetation is usually assessed by field surveys. Application of detailed sedimentary DNA (sedDNA) records has the potential to enhance our understanding of vegetation changes, but studies systematically investigating the power of this proxy are rare to date. This study compares sedDNA metabarcoding and pollen records from surface sediments of 31 lakes along a north–south gradient of increasing forest cover in northern Siberia (Taymyr peninsula) with data from field surveys in the surroundings of the lakes. sedDNA metabarcoding recorded 114 plant taxa, about half of them to species level, while pollen analyses identified 43 taxa, both exceeding the 31 taxa found by vegetation field surveys. Increasing Larix percentages from north to south were consistently recorded by all three methods and principal component analyses based on percentage data of vegetation surveys and DNA sequences separated tundra from forested sites. Comparisons of the ordinations using procrustes and protest analyses show a significant fit among all compared pairs of records. Despite similarities of sedDNA and pollen records, certain idiosyncrasies, such as high percentages of Alnus and Betula in all pollen and high percentages of Salix in all sedDNA spectra, are observable. Our results from the tundra to single‐tree tundra transition zone show that sedDNA analyses perform better than pollen in recording site‐specific richness (i.e., presence/absence of taxa in the vicinity of the lake) and perform as well as pollen in tracing vegetation composition.     

四川螺髻山表土和化石孢粉揭示的环境生态意义

通过四川螺髻山（1984、2017年）表土和化石孢粉与植被的关系分析发现：孢粉组合中木本植物含量（91.0%）占绝对优势,松属、冷杉属、青冈属、常绿栎类、落叶栎类、桤木属、杜鹃花科、禾本科和蒿属为主要花粉类型;人工次生林花粉组合能很好地反映母体植被的群落特征,并可指示优势种的存在和人类活动痕迹;针阔混交林花粉组合能较好地反映群落整体特征,花粉类型能与母体植被中优势种较好地对应;常绿阔叶林、针叶林和灌丛草甸的花粉组合难以反映母体植被的群落特征。DCA表明,花粉谱的百分含量,能较好地区分人工扰动植被和天然植被,但人工次生林、常绿阔叶林和针阔混交林之间以及针叶林和灌丛草甸之间未能区分。30年前后表土孢粉组合变化明显,1984年样品以松属、青冈属和桤木属等花粉为主,而2017年采样分析中松属、桤木属花粉占绝对优势。随着人类活动的加强,部分地区松树和桤木大面积飞播或种植,表土孢粉组合对当地植被指示意义与扰动强度呈负相关关系。现生植被与表土孢粉组合的差异能为恢复古植被和古环境提供参考,但原生植被已经被破坏地区的孢粉图谱,就很难作为重建历史时期植被的根据。本研究可以为亚热带山地利用孢粉学恢复第四纪时期植被与气候,探讨人类活动与环境的关系提供理论依据和实践参考。     

Soil phosphorus as a control of productivity and openness in temperate interglacial forest ecosystems

Aim Observations of long chronosequences in forest ecosystems show that, after some millennia of build‐up, biomass declines in relation to the slow depletion of soil phosphorus. Plants that dominate during this period of soil impoverishment have specialized strategies for P acquisition, including ectomycorrhiza or root clusters. We use quantitative, pollen‐based reconstructions of regional vegetation in four Quaternary warm stages (Holocene, Eemian, Holsteinian, Harreskovian) to test whether inferred forest cover and productivity changes are consistent with long‐term modification of soil nutrient pools. Location Southern Scandinavia (Denmark, southern Sweden). Methods The REVEALS model was used to estimate regional vegetation abundances of 25 pollen‐type‐equivalent taxa from pollen records of large sedimentary basins in southernmost Scandinavia. Based on the estimated regional vegetation, we then calculated time‐series of Ellenberg indicator values for L (light), R (soil reaction) and N (a productivity proxy). We classified the vegetation records into distinct phases and compared these phases and the samples using hierarchical clustering and ordination. Results All three interglacials developed coniferous or mixed forests. However, pure deciduous forests were never reached during the Holsteinian, while pure coniferous forests never developed in the Holocene. Above‐ground productivity was inferred to be low initially, peaking in the first third of the warm stages and then slowly declining (except during the Holocene). Dominant trees of the post‐peak phases all had ectomycorrhiza as a strategy for P acquisition, indicating that easily accessible P pools had become depleted. Increases in fire regimes may have amplified the inferred final drop in productivity. Mid/late Holocene productivity changes were much influenced by agricultural activities. Main conclusions REVEALS vegetation estimates combined with Ellenberg indicator values suggest a consistent pattern in warm stages of initially rising productivity, followed by a long and slow decline. The P‐acquisition strategies of dominant trees indicate that the decline reflects increasing P depletion of soils.     

Global ecosystems and fire: Multi‐model assessment of fire‐induced tree‐cover and carbon storage reduction

In this study, we use simulations from seven global vegetation models to provide the first multi‐model estimate of fire impacts on global tree cover and the carbon cycle under current climate and anthropogenic land use conditions, averaged for the years 2001–2012. Fire globally reduces the tree covered area and vegetation carbon storage by 10%. Regionally, the effects are much stronger, up to 20% for certain latitudinal bands, and 17% in savanna regions. Global fire effects on total carbon storage and carbon turnover times are lower with the effect on gross primary productivity (GPP) close to 0. We find the strongest impacts of fire in savanna regions. Climatic conditions in regions with the highest burned area differ from regions with highest absolute fire impact, which are characterized by higher precipitation. Our estimates of fire‐induced vegetation change are lower than previous studies. We attribute these differences to different definitions of vegetation change and effects of anthropogenic land use, which were not considered in previous studies and decreases the impact of fire on tree cover. Accounting for fires significantly improves the spatial patterns of simulated tree cover, which demonstrates the need to represent fire in dynamic vegetation models. Based upon comparisons between models and observations, process understanding and representation in models, we assess a higher confidence in the fire impact on tree cover and vegetation carbon compared to GPP, total carbon storage and turnover times. We have higher confidence in the spatial patterns compared to the global totals of the simulated fire impact. As we used an ensemble of state‐of‐the‐art fire models, including effects of land use and the ensemble median or mean compares better to observational datasets than any individual model, we consider the here presented results to be the current best estimate of global fire effects on ecosystems.     

Land‐use change outweighs projected effects of changing rainfall on tree cover in sub‐Saharan Africa

Global change will likely affect savanna and forest structure and distributions, with implications for diversity within both biomes. Few studies have examined the impacts of both expected precipitation and land use changes on vegetation structure in the future, despite their likely severity. Here, we modeled tree cover in sub‐Saharan Africa, as a proxy for vegetation structure and land cover change, using climatic, edaphic, and anthropic data (R² = 0.97). Projected tree cover for the year 2070, simulated using scenarios that include climate and land use projections, generally decreased, both in forest and savanna, although the directionality of changes varied locally. The main driver of tree cover changes was land use change; the effects of precipitation change were minor by comparison. Interestingly, carbon emissions mitigation via increasing biofuels production resulted in decreases in tree cover, more severe than scenarios with more intense precipitation change, especially within savannas. Evaluation of tree cover change against protected area extent at the WWF Ecoregion scale suggested areas of high biodiversity and ecosystem services concern. Those forests most vulnerable to large decreases in tree cover were also highly protected, potentially buffering the effects of global change. Meanwhile, savannas, especially where they immediately bordered forests (e.g. West and Central Africa), were characterized by a dearth of protected areas, making them highly vulnerable. Savanna must become an explicit policy priority in the face of climate and land use change if conservation and livelihoods are to remain viable into the next century.     

Interpretation of the last‐glacial vegetation of eastern‐central Europe using modern analogues from southern Siberia

Aim Interpretation of fossil pollen assemblages may benefit greatly from comparisons with modern palynological and vegetation analogues. To interpret the full‐ and late‐glacial vegetation in eastern‐central Europe we compared fossil pollen assemblages from this region with modern pollen assemblages from various vegetation types in southern Siberia, which presumably include the closest modern analogues of the last‐glacial vegetation of central Europe. Location Czech and Slovak Republics (fossil pollen assemblages); Western Sayan Mountains, southern Siberia (modern pollen assemblages). Methods Eighty‐eight modern pollen spectra were sampled in 14 vegetation types of Siberian forest, tundra and steppe, and compared with the last‐glacial pollen spectra from seven central European localities using principal components analysis. Results Both full‐ and late‐glacial pollen spectra from the valleys of the Western Carpathians (altitudes 350–610 m) are similar to modern pollen spectra from southern Siberian taiga, hemiboreal forest and dwarf‐birch tundra. The full‐glacial and early late‐glacial pollen spectra from lowland river valleys in the Bohemian Massif (altitudes 185–190 m) also indicate the presence of patches of hemiboreal forest or taiga. Other late‐glacial pollen spectra from the Bohemian Massif suggest an open landscape with steppe or tundra or a mosaic of both, possibly with small patches of hemiboreal forest. Main conclusions Our results are consistent with the hypothesis that during the full glacial and late glacial, the mountain valleys of the north‐western Carpathians supported taiga or hemiboreal forest dominated by Larix, Pinus cembra, Pinus sylvestris and Picea, along with some steppic or tundra formations. Forests tended to be increasingly open or patchy towards the west (Moravian lowlands), gradually passing into the generally treeless landscape of Bohemia, with possible woodland patches in locally favourable sites.     

Share the wealth: Trees with greater ectomycorrhizal species overlap share more carbon

The mutualistic symbiosis between forest trees and ectomycorrhizal fungi (EMF) is among the most ubiquitous and successful interactions in terrestrial ecosystems. Specific species of EMF are known to colonize specific tree species, benefitting from their carbon source, and in turn, improving their access to soil water and nutrients. EMF also form extensive mycelial networks that can link multiple root‐tips of different trees. Yet the number of tree species connected by such mycelial networks, and the traffic of material across them, are just now under study. Recently we reported substantial belowground carbon transfer between Picea, Pinus, Larix and Fagus trees in a mature forest. Here, we analyze the EMF community of these same individual trees and identify the most likely taxa responsible for the observed carbon transfer. Among the nearly 1,200 EMF root‐tips examined, 50%–70% belong to operational taxonomic units (OTUs) that were associated with three or four tree host species, and 90% of all OTUs were associated with at least two tree species. Sporocarp ¹³C signals indicated that carbon originating from labelled Picea trees was transferred among trees through EMF networks. Interestingly, phylogenetically more closely related tree species exhibited more similar EMF communities and exchanged more carbon. Our results show that belowground carbon transfer is well orchestrated by the evolution of EMFs and tree symbiosis.