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1 Disturbance may cause community composition across sites to become more or less homogenous, depending on the importance of different processes involved in community assembly. In north-eastern North America and Europe local (alpha) diversity of forest plants is lower in forests growing on former agricultural fields (recent forests) than in older (ancient) forests, but little is known about the influence of land-use history on the degree of compositional differentiation among sites (beta diversity). 2 Here we analyse data from 1446 sites in ancient and recent forests across 11 different landscapes in north-eastern North America and Europe to demonstrate decreases in beta diversity and in the strength of species-environment relationships in recent vs. ancient forests. 3 The magnitude of environmental variability among sites did not differ between the two forest types. This suggests the difference in beta diversity between ancient and recent forests was not due to different degrees of environmental heterogeneity, but rather to dispersal filters that constrain the pool of species initially colonizing recent forests. 4 The observed effects of community homogenization and weakened relationships between species distributions and environmental gradients appear to persist for decades or longer. The legacy of human land-use history in spatial patterns of biodiversity may endure, both within individual sites and across sites, for decades if not centuries.

QUESTIONS: (i) Is plant species richness related to patch configuration, quality, heterogeneity and history in very small forest patches? (ii) Is there a similar effect of patch configuration, quality, heterogeneity and history on all plant species or does it vary for different plant groups? (iii) Does the importance of patch configuration, quality, heterogeneity and history vary for plant species richness between patch size classes? LOCATION: Prignitz region, western part of Brandenburg, Germany METHODS: Plant species lists for 183 forest patches (0.08–12.7 ha) were compiled. We conducted a hierarchical partitioning analysis to test which of the four patch variables – configuration, quality, heterogeneity and history – explained most of the variance in plant species richness and richness of various plant groups. RESULTS: Irrespective of patch size class, species richness is mainly predicted by patch quality and heterogeneity. In particular, generalists are primarily affected by the soil moisture and nutrient level, and specialists additionally by patch heterogeneity and history. A higher historical habitat quality, together with long habitat continuity, led to an increase in forest specialists and geophytes in all patches. In the >5‐ha patches only historical habitat quality accounted for a significant increase in the number of species with short‐distance dispersal capacity in all species. Regardless of patch size, the number of locally infrequent species increased significantly with historical habitat quality. The effect of patch configuration was negligible, except for space‐filling trees and shrubs, when considering all patches. CONCLUSIONS: Our more comprehensive statistical approach shows, in general, that all complex variables have an effect on species richness and should be considered in future studies. The fact that patch heterogeneity and quality are the main predictors of species richness indicates that very small forest patches covered a range of highly structured patches and sites having various growth conditions. We assume that the century‐old tradition of private ownership with smallholder social structure is the main reason for variability in stand structure and species assemblage, resulting in the uniqueness of each patch. Since the number of locally infrequent plant species increases significantly with historical habitat quality, we suggest that it is particularly important to consider this variable in future studies on the decline of infrequent plant species.

Background Multi-purpose use of forests in a sustainable way forces a recognition of how introduction of alien woody species in forests with different land use histories affect native plants other than trees. Lingonberry Vaccinium vitis-idaea is an important understory component of temperate and boreal forests and provider of valuable non-wood forest products. Here we studied effects of land use changes and introduction of Northern red oak Quercus rubra on lingonberry in mesic Scots pine forests (in central Poland). We measured lingonberry cover, height of shoots, biomass of stems and leaves, and fruit productivity. Shoots were collected within 200 research plots located in recent and ancient Scots pine forests, with and without Q. rubra . Results We found that V. vitis-idaea reached lower cover, aboveground biomass and fruit production in recent than ancient forests and in forests with than without Q. rubra . The fruit production in recent pine forest was only 2% of that reported in ancient pine forest, and V. vitis-idaea did not reproduce generatively in forests with Q. rubra . Biomass and carbon sequestration of V. vitis-idaea in forests with alien (invasive) trees decreased by 75% compared to ancient pine forest. Effects were also clear at the individual shoot level – in less suitable conditions we found taller heights and higher biomass allocation into stems than foliage. Biomass allocation in fruiting and non-fruiting shoots in pine forests was also different – less of the dry biomass of fruiting shoots was allocated to leaves than to stems. Conclusions In the age of high interest in ecosystem services and discussions about usage of alien tree species as alternatives in forest management, our results clearly indicate disruption of ecosystem services provided by V. vitis-idaea in the presence of Q. rubra . Lingonberry benefited from the continuity of forest land use, however, regardless of land-use legacy, alien tree introduction led to decline in abundance of species crucial for ecosystem functioning. Therefore, to maintain valuable native species and for conservation of ecosystem services delivery, we suggest limiting the introduction of Q. rubra in areas with abundant V. vitis-idaea , especially in forests with continuous forest land-use history.

Questions: At least half of the European woodland is recent and most stands have been under heavy management since the Industrial Revolution. Woodland herbs have adapted to the conditions of the forest interior and many species occur at lower frequencies in recent habitats as well as in young stands. We compared herbaceous plant composition of forests with different land-use history and looked for life-history traits that could be responsible for the differences. Location: Bakony Mountains, Western Hungary, Central Europe. Methods: Adjacent old ancient, young ancient and post-agricultural recent forest stands were surveyed in the submontane beech zone in Western Hungary. Frequency data were recorded for herbaceous plants, indicator species analysis was carried out and trait-based species groups were identified. Results: Species composition differed among all three categories. The most profound difference was in the occurrences of ancient forest species (species with low dispersal ability and early flowering). These were most abundant in old ancient forests, while three species were entirely missing from recent stands. Young stands were dominated by few shade-tolerant herbs and lacked most ancient forest species. Post-agricultural woodland was largely dominated by competitive species. Conclusions: Current forest management as well as past agricultural use influenced herb composition. Reintroduction of ancient forest herbs might be necessary in the recent forests.

Revealing the interactive effects of multiple environmental change drivers (water deficits, nitrogen (N) deposition, land-use change) is crucial for evaluating actual and possible future changes in forest ecosystem functioning. Here, we analyse whether and to what extent combined effects of spring and summer water deficits and variable amounts of N deposition affect radial growth of beech trees growing on forest sites with a different forest history. Dendrochronological data showed that trees growing on ancient forest sites (forest continuity > 200 years) exhibit a higher negative growth response under high N deposition and simultaneous spring water deficits than trees growing on recent (post-agricultural) forest sites. Based on additional analyses of the fine root system and masting behaviour, we propose two different mechanisms to explain differing influences of N deposition and water deficits on negative radial growth responses in recent and ancient forests: (1) for both forest history types, growth reductions during summer water deficits result from the antagonistic effects of elevated N deposition according to the ‘resource optimization hypothesis’. The tendency towards higher negative growth responses in recent forests seems to be caused by a higher fine root mortality and lower standing fine root biomass compared to ancient forests; (2) higher growth reductions in ancient forests during spring water deficits are likely the result of mass fructification, which is enhanced by N deposition. We conclude that nutrient cycling may differ between forests with contrasting forest history, which can modulate the growth trajectories of forests in response to multiple, co-occurring environmental changes.

Agricultural expansion has resulted in loss of natural habitats worldwide, threatening remnant populations and communities. However, during the past several centuries in much of Europe and eastern North America, extensive forest clearance (usually resulting in <20% forest cover) has been followed by widespread expansion of forest onto abandoned agricultural land. Since ancient forests (those remaining at the peak of forest loss) provide seed sources for recolonization of recent (post-agricultural) forests, the extent of forest loss should influence the rate of population recovery in recent forest. Data compiled from the literature for 10 regions of Europe and North America indicate that the proportion of ancient forest in a landscape can account for >65% of the variation in species diversity of \"ancient forest herbs\" in recent forests relative to ancient forests. This result suggests that ancient-forest loss has had a strong influence on the recovery of herbaceous plant diversity in post-agricultural forests. In addition, a simple mainland-island metapopulation model suggests that with increasing severity of habitat loss the return time to equilibrium patch occupancy in recent forests is greatly extended, perhaps for centuries after forest clearance has ceased and restoration begun.

Question: (i) How does former land use and land use intensity affect seed bank development during post-agricultural succession? (ii) How does time since the last clear-cut change seed bank composition during post-clear-cut succession? Methods: One data set was compiled per succession type using the following selection criteria: (i) the data set included a successional series, (ii) plots were located in mesotrophic forest plant communities and (iii) vegetation data were available. The post-agricultural succession data set comprised 76 recent forest plots (eight studies); the post-clear-cut succession data set comprised 218 ancient forest plots (three studies). Each data set was analysed separately using either linear mixed models or generalized linear models, controlling for both environmental heterogeneity and variation between study locations. Results: In the post-agricultural succession data set, land use and time significantly affected nearly all the studied seed bank characteristics. Seed banks on former arable land recovered poorly even after 150 year of restored forest cover, whereas moderate land use intensities (grasslands, heathlands) yielded more rapid seed bank recovery. Time was a significant determinant of all but two soil seed bank characteristics during post-clear-cut succession. Seed banks in managed ancient forest differed strongly in their characteristics compared to primary forest seed banks. Conclusions: Forest seed banks bear the marks of former land use and/or forest management and continue to do so for at least 150 years. Nevertheless, time since the last major disturbance, being either former land use or clear-cutting, remains a significant determinant of the seed bank.

Changes in land use have been shown to have profound effects on forest plant community structure and diversity. Dispersal limitation has been invoked as a major factor hampering colonization of forest plant species, while seed-sowing experiments and performance observations have provided some evidence for recruitment limitation determining forest plant distribution in post-agricultural forests. However, most of these studies were relatively short-term, and very few studies have investigated long-term growth rates of populations occurring in recent and ancient forests. In this study, matrix models using demographic data collected for four consecutive years were used to study the effect of forest age on population dynamics of the temperate forest herb Primula elatior. A life table response experiment (LTRE) and elasticity analysis were used to analyze the effect of forest age on population growth rate (Λ) and to decompose the effect of forest age on Λ into contributions from each matrix element. Population growth increased logarithmically with increasing forest age. Bootstrap analyses showed that populations located in very recent forests (<50 years old) had growth rates that were significantly <1, whereas populations located in forests > 150 years old had growth rates that were significantly >1. Summed elasticities for individual growth significantly decreased with increasing forest age, whereas summed elasticities for survival and fertility significantly increased with increasing forest age. The LTRE analysis showed that the increase in Λ with increasing forest age was mainly due to increased seedling and juvenile growth and increased juvenile and adult survival. Our results indicate that past agricultural land use has long-lasting effects on the demography of forest herbs and may provide an additional mechanistic explanation for the poor colonization capacity of many forest herbs in post-agricultural forests.

Historical ecology gives a reference point to explain the contemporary state of particular ecosystems as well as entire landscapes. In this study we examined the quantitative changes in forest cover in the central part of the Sudety Massif (area ca. 1,120 [km.sup.2]) during the last 250 years. The information regarding forest patch distribution and its changes was derived by comparison of maps from 1747 and the 1970s drawn at scales of 1:33,000 and 1:25,000, respectively. To examine the effect of environmental variables (topography and soil conditions) and human population density on forest patch distribution and its changes (afforestation, deforestation), a set of 100 circular plots with a diameter of 1 km was established. The influence of explanatory variables was examined using regression tree methods. Changes at the level of the entire landscape were tested using a set of 25 landscape windows (5x5 km each). We found that the overall forest cover increased to 36.4% in the twentieth century from 30.4% in the middle of the eighteenth century. The ancient forests constituted 59% of the total forest area existing more recently. The forests in the eighteenth century occurred mostly on steep slopes, deep valley bottoms, and summits. The land relief explains more than half of the total variation in forest distribution ([R.sup.2] = 0.56). The effects of soil type and human population density were negligible. The contemporary forest pattern results from both land relief and the historical pattern of human population density in the middle of the eighteenth century ([R.sup.2] = 0.64), while the effect of soil type was negligible. The pattern of deforestation ([R.sup.2] = 0.53) and afforestation ([R.sup.2] = 0.36) results from both land relief as well as recent and nineteenth-century human population density. About 83% of the recent forest area is in physical contact with patches of the ancient forest, which provides an optimistic outlook for the migration of ancient forest species into new areas. Furthermore, changes in landscape structure reveal increased connectivity among forest patches, with potential benefits for the migration of forest species with long-range dispersal. Keywords ancient forests; deforestation; historical ecology; landscape fragmentation; recent forests

In Europe, many contemporary forests are not continuous, but were established on former agricultural land in territories of deserted villages. We aimed to explore whether medieval settlement activities of only 60 years irreversibly changed soil properties and whether these changes were reflected by contemporary forest vegetation. The research was performed in the deserted village of Kří, which existed from 1357 to ca 1420 AD in a current oak forest on sandy soils in the Czech Republic. We identified four former land-use types (building sites, courtyards, the village square, and gardens), where we analyzed soil properties and plant species composition of the forest understory. Higher concentrations of plant-available P, K, Ca, and Mg as well as the total concentrations of organic C and trace elements (As, Cd, Cu, and Zn) were recorded at former building sites with neutral soils, compared to the other former land-use types with acidic soils. The four-times higher species richness at former building sites compared to the former gardens indicate the strong effect of soil pH and nutrient availability on plant species composition, even on the spatial scale of several square metres. Understanding recent patterns in soil properties and biodiversity in Central European forests requires detailed knowledge of former land use over centuries. Analysis of contemporary soil properties together with analysis of vegetation can help to identify former land-use types in deserted villages.