Long-term shift towards shady and nutrient-rich habitats in Central European temperate forests.

Biodiversity world-wide has been under increasing anthropogenic pressure in the past century. The long-term response of biotic communities has been tackled primarily by focusing on species richness, community composition and functionality. Equally important are shifts between entire communities and habitat types, which remain an unexplored level of biodiversity change. We have resurveyed > 2000 vegetation plots in temperate forests in central Europe to capture changes over an average of five decades. The plots were assigned to eight broad forest habitat types using an algorithmic classification system. We analysed transitions between the habitat types and interpreted the trend in terms of changes in environmental conditions. We identified a directional shift along the combined gradients of canopy openness and soil nutrients. Nutrient-poor open-canopy forest habitats have declined strongly in favour of fertile closed-canopy habitats. However, the shift was not uniform across the whole gradients. We conclude that the shifts in habitat types represent a century-long successional trend with significant consequences for forest biodiversity. Open forest habitats should be urgently targeted for plant diversity restoration through the implementation of active management. The approach presented here can be applied to other habitat types and at different spatio-temporal scales.

Evaluating plant lineage losses and gains in temperate forest understories: a phylogenetic perspective on climate change and nitrogen deposition.

Global change has accelerated local species extinctions and colonizations, often resulting in losses and gains of evolutionary lineages with unique features. Do these losses and gains occur randomly across the phylogeny? We quantified: temporal changes in plant phylogenetic diversity (PD); and the phylogenetic relatedness (PR) of lost and gained species in 2672 semi-permanent vegetation plots in European temperate forest understories resurveyed over an average period of 40 yr. Controlling for differences in species richness, PD increased slightly over time and across plots. Moreover, lost species within plots exhibited a higher degree of PR than gained species. This implies that gained species originated from a more diverse set of evolutionary lineages than lost species. Certain lineages also lost and gained more species than expected by chance, with Ericaceae, Fabaceae, and Orchidaceae experiencing losses and Amaranthaceae, Cyperaceae, and Rosaceae showing gains. Species losses and gains displayed no significant phylogenetic signal in response to changes in macroclimatic conditions and nitrogen deposition. As anthropogenic global change intensifies, temperate forest understories experience losses and gains in specific phylogenetic branches and ecological strategies, while the overall mean PD remains relatively stable.

Les changements globaux accélèrent les processus de colonisation et d'extinction locales d'espèces, aboutissant à des gains ou à des pertes de lignées évolutives uniques. Ces gains et pertes se produisent-ils de manière aléatoire dans l'arbre phylogénétique ? Nous avons mesuré: les changements de diversité phylogénétique; et la parenté phylogénétique des espèces végétales gagnées ou perdues dans 2672 placettes semi-permanentes disposées dans le sous-bois de forêts tempérées d'Europe sur une période moyenne de 40 ans. Une fois corrigée par la richesse spécifique, la diversité phylogénétique a légèrement augmenté au cours du temps dans les différentes placettes. Les espèces perdues ont une plus grande parenté phylogénétique que les espèces gagnées. Les espèces gagnées sont donc issues d'un plus grand nombre de lignées évolutives que les espèces perdues. Certaines lignées ont gagné ou perdu davantage d'espèces que ce qui est prédit par le hasard : les Ericaceae, les Fabaceae et les Orchidaceae ayant davantage perdu, tandis que les Amaranthaceae, les Cyperaceae, et les Rosaceae ont plus gagné. Il n'y a pas de signal phylogénétique des gains ou pertes d'espèces en réponse aux changements de conditions macroclimatiques ou des dépôts atmosphériques d'azote. Alors que les changements globaux d'origine anthropique s'intensifient, les sous-bois des forêts tempérées connaissent des gains et des pertes de certaines lignées évolutives et de certaines stratégies écologiques, sans que la diversité phylogénétique moyenne ne s'en trouve véritablement affectée.

El cambio global ha acelerado las extinciones y colonizaciones a escala local, lo que a menudo ha supuesto pérdidas y ganancias de linajes evolutivos con características únicas. Ahora bien, ¿estas pérdidas y ganancias ocurren aleatoriamente a lo largo de la filogenia? Cuantificamos: los cambios temporales en la diversidad filogenética de las plantas; y la relación filogenética de las especies perdidas y ganadas en 2.672 parcelas de vegetación semipermanente en sotobosques templados europeos y re-muestreadas durante un período promedio de 40 años. Al controlar por las diferencias en la riqueza de especies, la diversidad filogenética aumentó ligeramente con el tiempo y entre parcelas. Además, las especies perdidas dentro de las parcelas exhibieron un mayor grado de relación filogenética que las especies ganadas. Esto implica que las especies ganadas se originaron en un conjunto de linajes evolutivos más diversos que las especies perdidas. Ciertos linajes también perdieron y ganaron más especies de las esperadas aleatoriamente: Ericaceae, Fabaceae y Orchidaceae experimentaron pérdidas y Amaranthaceae, Cyperaceae y Rosaceae mostraron ganancias. Las pérdidas y ganancias de especies no mostraron ninguna señal filogenética significativa en respuesta a los cambios en las condiciones macro-climáticas y la deposición de nitrógeno. A medida que se intensifica el cambio global antropogénico, los sotobosques temperados experimentan pérdidas y ganancias en ramas filogenéticas y estrategias ecológicas específicas, mientras que la diversidad filogenética media general permanece relativamente estable.

Combining multiple investigative approaches to unravel functional responses to global change in the understorey of temperate forests.

Plant communities are being exposed to changing environmental conditions all around the globe, leading to alterations in plant diversity, community composition, and ecosystem functioning. For herbaceous understorey communities in temperate forests, responses to global change are postulated to be complex, due to the presence of a tree layer that modulates understorey responses to external pressures such as climate change and changes in atmospheric nitrogen deposition rates. Multiple investigative approaches have been put forward as tools to detect, quantify and predict understorey responses to these global-change drivers, including, among others, distributed resurvey studies and manipulative experiments. These investigative approaches are generally designed and reported upon in isolation, while integration across investigative approaches is rarely considered. In this study, we integrate three investigative approaches (two complementary resurvey approaches and one experimental approach) to investigate how climate warming and changes in nitrogen deposition affect the functional composition of the understorey and how functional responses in the understorey are modulated by canopy disturbance, that is, changes in overstorey canopy openness over time. Our resurvey data reveal that most changes in understorey functional characteristics represent responses to changes in canopy openness with shifts in macroclimate temperature and aerial nitrogen deposition playing secondary roles. Contrary to expectations, we found little evidence that these drivers interact. In addition, experimental findings deviated from the observational findings, suggesting that the forces driving understorey change at the regional scale differ from those driving change at the forest floor (i.e., the experimental treatments). Our study demonstrates that different approaches need to be integrated to acquire a full picture of how understorey communities respond to global change.

Directional turnover towards larger-ranged plants over time and across habitats.

Species turnover is ubiquitous. However, it remains unknown whether certain types of species are consistently gained or lost across different habitats. Here, we analysed the trajectories of 1827 plant species over time intervals of up to 78 years at 141 sites across mountain summits, forests, and lowland grasslands in Europe. We found, albeit with relatively small effect sizes, displacements of smaller- by larger-ranged species across habitats. Communities shifted in parallel towards more nutrient-demanding species, with species from nutrient-rich habitats having larger ranges. Because these species are typically strong competitors, declines of smaller-ranged species could reflect not only abiotic drivers of global change, but also biotic pressure from increased competition. The ubiquitous component of turnover based on species range size we found here may partially reconcile findings of no net loss in local diversity with global species loss, and link community-scale turnover to macroecological processes such as biotic homogenisation.

Environmental drivers interactively affect individual tree growth across temperate European forests.

Forecasting the growth of tree species to future environmental changes requires a better understanding of its determinants. Tree growth is known to respond to global-change drivers such as climate change or atmospheric deposition, as well as to local land-use drivers such as forest management. Yet, large geographical scale studies examining interactive growth responses to multiple global-change drivers are relatively scarce and rarely consider management effects. Here, we assessed the interactive effects of three global-change drivers (temperature, precipitation and nitrogen deposition) on individual tree growth of three study species (Quercus robur/petraea, Fagus sylvatica and Fraxinus excelsior). We sampled trees along spatial environmental gradients across Europe and accounted for the effects of management for Quercus. We collected increment cores from 267 trees distributed over 151 plots in 19 forest regions and characterized their neighbouring environment to take into account potentially confounding factors such as tree size, competition, soil conditions and elevation. We demonstrate that growth responds interactively to global-change drivers, with species-specific sensitivities to the combined factors. Simultaneously high levels of precipitation and deposition benefited Fraxinus, but negatively affected Quercus' growth, highlighting species-specific interactive tree growth responses to combined drivers. For Fagus, a stronger growth response to higher temperatures was found when precipitation was also higher, illustrating the potential negative effects of drought stress under warming for this species. Furthermore, we show that past forest management can modulate the effects of changing temperatures on Quercus' growth; individuals in plots with a coppicing history showed stronger growth responses to higher temperatures. Overall, our findings highlight how tree growth can be interactively determined by global-change drivers, and how these growth responses might be modulated by past forest management. By showing future growth changes for scenarios of environmental change, we stress the importance of considering multiple drivers, including past management and their interactions, when predicting tree growth.

Global environmental change effects on plant community composition trajectories depend upon management legacies.

The contemporary state of functional traits and species richness in plant communities depends on legacy effects of past disturbances. Whether temporal responses of community properties to current environmental changes are altered by such legacies is, however, unknown. We expect global environmental changes to interact with land-use legacies given different community trajectories initiated by prior management, and subsequent responses to altered resources and conditions. We tested this expectation for species richness and functional traits using 1814 survey-resurvey plot pairs of understorey communities from 40 European temperate forest datasets, syntheses of management transitions since the year 1800, and a trait database. We also examined how plant community indicators of resources and conditions changed in response to management legacies and environmental change. Community trajectories were clearly influenced by interactions between management legacies from over 200 years ago and environmental change. Importantly, higher rates of nitrogen deposition led to increased species richness and plant height in forests managed less intensively in 1800 (i.e., high forests), and to decreases in forests with a more intensive historical management in 1800 (i.e., coppiced forests). There was evidence that these declines in community variables in formerly coppiced forests were ameliorated by increased rates of temperature change between surveys. Responses were generally apparent regardless of sites' contemporary management classifications, although sometimes the management transition itself, rather than historic or contemporary management types, better explained understorey responses. Main effects of environmental change were rare, although higher rates of precipitation change increased plant height, accompanied by increases in fertility indicator values. Analysis of indicator values suggested the importance of directly characterising resources and conditions to better understand legacy and environmental change effects. Accounting for legacies of past disturbance can reconcile contradictory literature results and appears crucial to anticipating future responses to global environmental change.

Using historical ecology to reassess the conservation status of coniferous forests in Central Europe.

Forests cover approximately one-third of Central Europe. Oak (Quercus) and European beech (Fagus sylvatica) are considered the natural dominants at low and middle elevations, respectively. Many coniferous forests (especially of Picea abies) occur primarily at midelevations, but these are thought to have resulted from forestry plantations planted over the past 200 years. Nature conservation and forestry policy seek to promote broadleaved trees over conifers. However, there are discrepancies between conservation guidelines (included in Natura 2000) and historical and palaeoecological data with regard to the distribution of conifers. Our aim was to bring new evidence to the debate on the conservation of conifers versus broadleaved trees at midelevations in Central Europe. We created a vegetation and land-cover model based on pollen data for a highland area of 11,300 km2 in the Czech Republic and assessed tree species composition in the forests before the onset of modern forestry based on 18th-century archival sources. Conifers dominated the study region throughout the entire Holocene (approximately 40-60% of the area). Broadleaved trees were present in a much smaller area than envisaged by current ideas of natural vegetation. Rather than casting doubt on the principles of Central European nature conservation in general, our results highlight the necessity of detailed regional investigations and the importance of historical data in challenging established notions on the natural distribution of tree species.

Dynamics of herbaceous vegetation during four years of experimental coppice introduction.

Understanding the effects of coppicing on forest ecosystems is important for progress towards sustainable forest management. A newly established coppicing experiment in a secondary temperate deciduous forest in the SE Czech Republic provides a rather unique insight into succession driven by canopy thinning in a forest still lacking species typical for forests established since long time ago. Herbaceous layer vegetation was monitored for four subsequent years in 2012-2015. We focused on the influence of canopy thinning intensity in two different forest types defined by dominant tree species (oak and lime). Our results showed that the opening of the canopy had immediate effects on herbaceous vegetation. Coverage, species richness and compositional patterns followed the coppicing intensity gradient. The dominant tree species had contrasting effects. Under oak, the reaction to coppicing was weak. Under lime, strong reaction both related to coppicing intensity and temporal development was observed. Herbs with short life cycle had the greatest contribution, but perennial grasses also began to increase their coverage after coppicing. Several invasive species, mostly short-lived herbs, emerged but are supposed to retreat as the succession will proceed. We conclude that coppice introduction to a secondary forest led to contrasting patterns related to dominant tree species. The marked difference was probably due to the slow sucession towards a future forest community saturated by species. This process may be now further diversified by coppicing management.

Patterns of functional diversity of two trophic groups after canopy thinning in an abandoned coppice.

Coppice abandonment had negative consequences for biodiversity of forest vegetation and several groups of invertebrates. Most coppicing restoration studies have focused only on a single trophic level despite the fact that ecosystems are characterized by interactions between trophic levels represented by various groups of organisms. To address the patterns of functional diversity in the perspective of coppicing restoration, we studied the short-term effects of conservation-motivated tree canopy thinning in an abandoned coppice-with-standards in Central Europe, a region where such attempts have been rare so far. The functional diversity of vascular plants and spiders, chosen as two model trophic groups within a forest ecosystem, was compared between thinned and control forest patches. To characterize functional patterns, we examined several functional traits. These traits were assigned into two contrasting categories: response traits reflecting a change of environment (for both vascular plants and spiders) and effect traits influencing the ecosystem properties (only for vascular plants). Functional diversity was analysed by CCA using two measures: community-weighted means (CWM) and Rao's quadratic diversity (RaoQ). CCA models revealed that the canopy thinning had a positive effect on diversity of the response traits of both trophic groups and negatively influenced the diversity of effect traits. In addition, we found distinct seasonal dynamics in functional diversity of the spider communities, which was probably linked to leaf phenology of deciduous trees and therefore an effect trait not directly examined in this study. We conclude that canopy thinning affected functional diversity across trophic groups during the initial phase of coppicing restoration. With necessary precautions, careful canopy thinning can be effectively applied in the restoration of functional diversity in abandoned coppices.

The paradox of long-term ungulate impact: increase of plant species richness in a temperate forest.

QUESTIONS: Did high densities of wild ungulates cause a decline in plant species richness in a temperate oakwood? How did species composition change after nearly five decades? Did ungulates facilitate the spread of ruderal species and supress endangered species? Did dispersal strategies play a role in these processes? LOCATION: Krumlov Wood, SE Czech Republic. METHODS: In 2012, we resampled 58 quasi-permanent vegetation plots first surveyed in 1960s. Between the surveys, 36 plots were enclosed in a game preserve with artificially high density of ungulates (mostly deer, mouflon and wild boar; ca. 55 animals per square km). We analysed the differences in temporal changes between plots inside and outside the game preserve, focusing on species diversity and composition. We assessed species characteristics relevant to grazing to understand compositional changes. RESULTS: Ungulates significantly increased alpha and gamma diversity and caused significant vegetation homogenization inside the game preserve. Vegetation homogenization and the increase in species richness resulted from massive enrichment by ruderal species. However, richness of endangered species decreased. Species dispersed by animals internally (endozoochory) increased, while species dispersed externally (epizoochory) or by wind (anemochory) decreased. CONCLUSIONS: Contrary to our expectations, our long-term data showed that artificially high ungulate densities substantially increased plant species richness. Apparently, the establishment of ruderal herbs was supported by frequent disturbances and ungulate-mediated dispersal. At the same time, species richness of non-ruderal plants did not change, probably because ungulates hindered the regeneration of woody species and maintained an open forest canopy. In conclusion, high ungulate density led to the spread of ruderal species, which in turn strongly contributed to the observed shift towards nutrient-richer conditions and taxonomically more homogenous communities.

Trends and events through seven centuries: the history of a wetland landscape in the Czech Republic.

Environmental change can be viewed as the combined result of long-term processes and singular events. While long-term trends appear to be readily available for observation (in the form of temporal comparisons or space-for-time substitution), it is more difficult to gain information on singular events in the past, although these can be equally significant in shaping ecosystems. We examined the past 700 years in the history of a lowland wetland landscape in the Czech Republic with the help of palaeoecological, ecological, landscape archaeological, and archival data. Macrofossil and pollen data were compared to known drainage works in the area and historical climatological data. Trends and events in habitat conditions were assessed using species indicator values. Results showed that ecological succession was the general process in the study area, detected as a trend towards eutrophication, desiccation and vegetation closure. Short-term events influenced development at the sites mainly from the second half of the 19th century. This is consistent with drainage history, although bias related to sample frequency cannot be excluded. On the whole, long-term trends and discrete events were complementary on different scales. We conclude that humans facilitated and accelerated background processes, which can be most likely associated with the succession of open wetlands towards terrestrial ecosystems.

Combining community resurvey data to advance global change research.

More and more ecologists have started to resurvey communities sampled in earlier decades to determine long-term shifts in community composition and infer the likely drivers of the ecological changes observed. However, to assess the relative importance of, and interactions among, multiple drivers joint analyses of resurvey data from many regions spanning large environmental gradients are needed. In this paper we illustrate how combining resurvey data from multiple regions can increase the likelihood of driver-orthogonality within the design and show that repeatedly surveying across multiple regions provides higher representativeness and comprehensiveness, allowing us to answer more completely a broader range of questions. We provide general guidelines to aid implementation of multi-region resurvey databases. In so doing, we aim to encourage resurvey database development across other community types and biomes to advance global environmental change research.

Microclimate moderates plant responses to macroclimate warming.

Recent global warming is acting across marine, freshwater, and terrestrial ecosystems to favor species adapted to warmer conditions and/or reduce the abundance of cold-adapted organisms (i.e., "thermophilization" of communities). Lack of community responses to increased temperature, however, has also been reported for several taxa and regions, suggesting that "climatic lags" may be frequent. Here we show that microclimatic effects brought about by forest canopy closure can buffer biotic responses to macroclimate warming, thus explaining an apparent climatic lag. Using data from 1,409 vegetation plots in European and North American temperate forests, each surveyed at least twice over an interval of 12-67 y, we document significant thermophilization of ground-layer plant communities. These changes reflect concurrent declines in species adapted to cooler conditions and increases in species adapted to warmer conditions. However, thermophilization, particularly the increase of warm-adapted species, is attenuated in forests whose canopies have become denser, probably reflecting cooler growing-season ground temperatures via increased shading. As standing stocks of trees have increased in many temperate forests in recent decades, local microclimatic effects may commonly be moderating the impacts of macroclimate warming on forest understories. Conversely, increases in harvesting woody biomass--e.g., for bioenergy--may open forest canopies and accelerate thermophilization of temperate forest biodiversity.

Resurveying historical vegetation data - opportunities and challenges.

BACKGROUND: Resurveying historical vegetation plots has become more and more popular in recent years as it provides a unique opportunity to estimate vegetation and environmental changes over the past decades. Most historical plots, however, are not permanently marked and uncertainty in plot location, in addition to observer bias and seasonal bias, may add significant error to temporal change. These errors may have major implications for the reliability of studies on long-term environmental change and deserve closer attention of vegetation ecologists. MATERIAL & METHODS: Vegetation data obtained from the resurveying of non-permanently marked plots are assessed for their potential to study environmental-change effects on plant communities and the challenges the use of such data have to meet. We describe the properties of vegetation resurveys distinguishing basic types of plots according to relocation error, and we highlight the potential of such data types for studying vegetation dynamics and their drivers. Finally, we summarise the challenges and limitations of resurveying non-permanently marked vegetation plots for different purposes in environmental change research. RESULTS AND CONCLUSIONS: Resampling error is caused by three main independent sources of error: error caused by plot relocation, observer bias, and seasonality bias. For relocation error, vegetation plots can be divided into permanent and non-permanent plots, while the latter are further divided into quasi-permanent (with approximate relocation) and non-traceable (with random relocation within a sampled area) plots. To reduce the inherent sources of error in resurvey data, the following precautions should be followed: (i) resurvey historical vegetation plots whose approximate plot location within a study area is known; (ii) consider all information available from historical studies in order to keep plot relocation errors low; (iii) resurvey at times of the year when vegetation development is comparable to the historical survey to control for seasonal variability in vegetation; (iv) keep a high level of experience of the observers to keep observer bias low; and (v) edit and standardise datasets before analyses.

Drivers of temporal changes in temperate forest plant diversity vary across spatial scales.

Global biodiversity is affected by numerous environmental drivers. Yet, the extent to which global environmental changes contribute to changes in local diversity is poorly understood. We investigated biodiversity changes in a meta-analysis of 39 resurvey studies in European temperate forests (3988 vegetation records in total, 17-75 years between the two surveys) by assessing the importance of (i) coarse-resolution (i.e., among sites) vs. fine-resolution (i.e., within sites) environmental differences and (ii) changing environmental conditions between surveys. Our results clarify the mechanisms underlying the direction and magnitude of local-scale biodiversity changes. While not detecting any net local diversity loss, we observed considerable among-site variation, partly explained by temporal changes in light availability (a local driver) and density of large herbivores (a regional driver). Furthermore, strong evidence was found that presurvey levels of nitrogen deposition determined subsequent diversity changes. We conclude that models forecasting future biodiversity changes should consider coarse-resolution environmental changes, account for differences in baseline environmental conditions and for local changes in fine-resolution environmental conditions.

Expansion to abandoned agricultural land forms an integral part of Silver fir dynamics.

Silver fir (Abies alba Mill.) is a tree species distributed mainly in central Europe. It once was a dominant tree species within some forests of this region. The causes for its rapid decline in the past two centuries have not yet been sufficiently explained. It is argued that human activities have been largely responsible for expansions and contractions of silver fir populations. On the basis of the current distribution of silver fir, historical maps and palaeoecological data, we describe the expansion of silver fir forests. We use fine resolution at the landscape level, an approach that has so far been neglected. Our study areas lies in the northern part of the White Carpathian Mountains, Czech Republic. The area comprises 7,045 ha, 65% of which is covered by forests. This landscape was shaped by early modern colonization from the 16th century onwards and has changed greatly since the decline of its traditional utilisation in the 19th and 20th centuries. The area of forests almost doubled from 1838 to 2005 while the area of pastures and arable land decreased. We identified 172 hectares of silver fir forests by field mapping, which represent 2.5% of the whole study area and 3.8% of its forested part. We used land use history variables (based on subsequent land cover maps from 1838, 1882 and 1956) and terrain variables (derived from a digital elevation model) in a logistic regression to model the probability of silver fir forest occurrence. Land use history was highly significantly correlated with the occurrence of silver fir forests. Approximately 59% of silver fir forests occur on land used as pastures in 1838, 28% are on former arable land, meadows and fallows, while only 13% have been forested continuously since the 19th century. We know from historical sources that the surrounding forests (now mainly Norway spruce monocultures) were dominated by silver fir up to the 1860s. Silver fir can act as a pioneer species. It can invade former agricultural land, which probably ensures the survival and periodical expansion of silver fir-dominated forests. Although silver fir has been thought to decline under human pressure, we suggest that the opposite may occur at the landscape level.

Coppice abandonment and its implications for species diversity in forest vegetation.

Coppicing, once a common type of management in European broadleaved forests, was abandoned in many places after WWII. This form of management provided a variety of structural and microclimatic conditions for tree and understorey vegetation. After the abandonment of this intensive management, succession towards mature close forests ensued, and suitable habitats for species ecologically connected to coppicing were reduced. In our study, we chose a region in central Europe where coppicing was the dominant type of forest management until the first half of the 20th century but was abandoned after WWII. We investigated long-term changes in both woody and herbaceous species composition in the Lower Morava UNESCO Biosphere Reserve using historical sources and vegetation plot resurveys from the 17th to the 21st century. The impact of coppice abandonment on vegetation composition and on the conservation value of forests was evaluated. Dominant tree species appeared to be very stable throughout the past four centuries, but changes occurred in their proportions. A shift from species rich oak-hornbeam woodland towards species poorer communities with increasing proportions of lime, ash and maple was observed after the abandonment of coppicing. The observed tendencies partly differed according to site and data source. The conservation value of forests was measured as the occurrence of red-list species, which were considerably reduced after coppice abandonment. To stop the process of biodiversity loss and support the goals of nature conservation, the re-establishment of coppice management is proposed.

Effects of simulated historical tree litter raking on the understorey vegetation in a central European forest.

QUESTION: What is the impact of simulated historical tree litter removal on understorey plants and soil properties in a temperate deciduous forest? What is the role of seasonal timing of tree litter removal on understorey plants? LOCATION: Podyjí National Park, Czech Republic. METHODS: We conducted an experiment in a randomized complete block design of 45 plots (5 × 5 m). Each block (N = 15) consisted of one plot for each of the three treatments. Treatments consisted of (i) tree litter removal during spring, (ii) tree litter removal during autumn, or (iii) no litter removal as control treatment. These treatments were repeated for a duration of four years. In each plot we recorded the understorey plant species composition and collected soil samples prior to treatment (year 0) and in each subsequent year (years 1-4). Temporal trends in species richness were analysed using repeated measures ANOVAs. The impact of the treatments on vegetation composition over time was analysed using Principal Response Curves. RESULTS: Total species richness per plot significantly changed over time, but this was not related to treatment. Annual species richness increased significantly, but only for the autumn treatment. Annual species also showed the highest inter-annual variation. Endangered species were not affected. When compared to the control treatment, the effect of autumn raking on species composition was stronger than the effect of spring raking. Although the amount of removed nutrients substantially exceeded ambient nitrogen input, no changes in soil conditions were detected. CONCLUSIONS: The season in which tree litter removal took place had a small but significant impact on the understorey vegetation, in particular affecting the germination and establishment of annual species. The large inter-annual variation in species richness calls for a long-term field experiment. The removal of nutrients via litter raking greatly exceeds atmospheric nutrient deposition, warranting a further investigation of litter raking as a potential tool for forest conservation.

Plant movements and climate warming: intraspecific variation in growth responses to nonlocal soils.

Most range shift predictions focus on the dispersal phase of the colonization process. Because moving populations experience increasingly dissimilar nonclimatic environmental conditions as they track climate warming, it is also critical to test how individuals originating from contrasting thermal environments can establish in nonlocal sites. We assess the intraspecific variation in growth responses to nonlocal soils by planting a widespread grass of deciduous forests (Milium effusum) into an experimental common garden using combinations of seeds and soil sampled in 22 sites across its distributional range, and reflecting movement scenarios of up to 1600 km. Furthermore, to determine temperature and forest-structural effects, the plants and soils were experimentally warmed and shaded. We found significantly positive effects of the difference between the temperature of the sites of seed and soil collection on growth and seedling emergence rates. Migrant plants might thus encounter increasingly favourable soil conditions while tracking the isotherms towards currently 'colder' soils. These effects persisted under experimental warming. Rising temperatures and light availability generally enhanced plant performance. Our results suggest that abiotic and biotic soil characteristics can shape climate change-driven plant movements by affecting growth of nonlocal migrants, a mechanism which should be integrated into predictions of future range shifts.

The Rise and Fall of Traditional Forest Management in Southern Moravia: A History of the Past 700 Years.

European broadleaved forests have been influenced by humans for centuries. Historical management practices are related to environmental conditions but the role of socio-economic factors is also important. For the successful restoration of traditional management for conservation purposes, detailed knowledge on management history and on the driving forces of historical forest changes is necessary. In order to reconstruct long-term spatio-temporal dynamics in forest management, we chose the Pálava Protected Landscape Area, Czech Republic and analyzed archival sources spanning the past seven centuries. Forests in the study area comprise two relatively large woods (Devín and Milovice) with different environmental conditions. Historical forest management in both woods was coppicing. The coppice cycle was lengthened from 7 years (14th century) to more than 30 years (19th century) with a fluctuating density of standards. After WWII, coppicing was completely abandoned. This led to pronounced changes in forest age structure accompanied by stand unification indicated by a sharp decrease in the Shannon index of age diversity. To study local attributes responsible for spatial patterns in coppice abandonment, we constructed a regression model with the date of abandonment as a dependent variable and three groups of explanatory variables: i) remoteness of forest parcels, (ii) morphometric environmental factors and iii) site productivity. In Devín Wood, coppicing was abandoned gradually with the pattern of abandonment related significantly to slope steepness and forest productivity. Poorly accessible upper slopes and low productive forest sites were abandoned earlier. By contrast, in Milovice Wood, where no clear topographic gradient is present, the abandonment of coppicing was not related to any of the variables we studied. Our study brings insights into the history and consequences of past management practices, and can be used in current attempts to re-establish coppice management for conservation purposes and as a source of sustainable energy.