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Context Plant populations in agricultural landscapes are mostly fragmented and their functional connectivity often depends on seed and pollen dispersal by animals. However, little is known about how the interactions of seed and pollen dispersers with the agricultural matrix translate into gene flow among plant populations. Objectives We aimed to identify effects of the landscape structure on the genetic diversity within, and the genetic differentiation among, spatially isolated populations of three temperate forest herbs. We asked, whether different arable crops have different effects, and whether the orientation of linear landscape elements relative to the gene dispersal direction matters. Methods We analysed the species’ population genetic structures in seven agricultural landscapes across temperate Europe using microsatellite markers. These were modelled as a function of landscape composition and configuration, which we quantified in buffer zones around, and in rectangular landscape strips between, plant populations. Results Landscape effects were diverse and often contrasting between species, reflecting their association with different pollen- or seed dispersal vectors. Differentiating crop types rather than lumping them together yielded higher proportions of explained variation. Some linear landscape elements had both a channelling and hampering effect on gene flow, depending on their orientation. Conclusions Landscape structure is a more important determinant of the species’ population genetic structure than habitat loss and fragmentation per se. Landscape planning with the aim to enhance the functional connectivity among spatially isolated plant populations should consider that even species of the same ecological guild might show distinct responses to the landscape structure.

The influence of environmental factors on the distribution and persistence of African elephants (Loxodonta africana) is pertinent to policy makers and managers to formulate balanced plans for different land‐use types.The study focuses on movement of elephants and how they utilize foraging areas in Sioma Ngwezi landscape in Zambia by answering the following questions: (1) Which environmental variables and land‐cover class predict the movement of elephants during the wet season in Sioma Ngwezi landscape? (2) What is the wet season suitable habitat for elephants in Sioma Ngwezi landscape? (3) What are the major wet season movement corridors for elephants in Sioma Ngwezi landscape?We used GPS telemetry data from the collared elephants to assess habitat connectivity. Maximum entropy (MaxEnt) and linkage mapper were the tools used to predict habitat suitability, movement corridors, and barriers in the landscape during the wet season.The study identified elevation, land cover, and NDVI as the most important environmental predictors that modify the dispersal of elephants in the landscape during the wet season. Additionally, a total of 36 potential wet season corridors were identified connecting 15 core areas mainly used for foraging and protection from poachers in the landscape. Of these, 24 corridors were highly utilized and are suggested as priority corridors for elephant movement in the landscape.The identified wet season habitats and functional corridors may help to combat elephant poaching by patrolling areas with high relative probability of elephant presence. The findings may also help abate human–elephant conflict such as crop‐raiding by managing identified corridors that run into agriculture zones in the game management area., The study focuses on movement of elephants and how they utilize foraging areas in wet season in Sioma Ngwezi landscape in Zambia.

In many agricultural landscapes, it is important to restore networks of forests to provide habitat and stepping stones for forest specialist taxa. More knowledge is, however, needed on how to facilitate the immigration of such taxa in restored forest patches. Here, we present the first chronosequence study to quantify the dynamics of immigration credits of forest specialist plants in post-arable forest patches. We studied the distribution of herbaceous forest specialist plant species in 54 post-arable broadleaved forest patches along gradients of age (20-140 years since forest establishment), distance from ancient forest (0-2,600 m) and patch area (0.5-9.6 ha). With linear mixed models, we estimated the effects of these factors on species richness, patch means of four dispersal-related plant traits and with generalized linear models on the occurrence of 20 individual species. Post-arable forest patch age and spatial isolation from ancient forest, but not patch size, were important predictors for species richness of forest specialists, suggesting that also small patches are valuable for habitat connectivity. Compared to species richness in ancient forest stands, the immigration credit was reduced by more than 90% after 80 years in post-arable forest patches contiguous to ancient forest compared to 40% after 80 years and 60% after 140 years in isolated patches (at least 100 m to next forest). Tall-growing species with adaptations to long-distance dispersal were faster colonizers, whereas species with heavy diaspores and clonal growth were slower to colonize. Synthesis and applications. We show that post-arable oak plantations have a high potential for restoration of forest herb vegetation. Dispersal-related plant traits play a key role in explaining interspecific differences among forest specialists. To facilitate forest herb immigration across all functional groups in agricultural landscapes, we suggest to create clusters of relatively small new forest patches nearby older forest with source populations.

Context Evidence for effects of habitat loss and fragmentation on the viability of temperate forest herb populations in agricultural landscapes is so far based on population genetic studies of single species in single landscapes. However, forest herbs differ in their life histories, and landscapes have different environments, structures and histories, making generalizations difficult. Objectives We compare the response of three slow-colonizing forest herbs to habitat loss and fragmentation and set this in relation to differences in life-history traits, in particular their mating system and associated pollinators. Methods We analysed the herbs’ landscape-scale population genetic structure based on microsatellite markers from replicate forest fragments across seven European agricultural landscapes. Results All species responded to reductions in population size with a decrease in allelic richness and an increase in genetic differentiation among populations. Genetic differentiation also increased with enhanced spatial isolation. In addition, each species showed unique responses. Heterozygosity in the self-compatible Oxalis acetosella was reduced in smaller populations. The genetic diversity of Anemone nemorosa, whose main pollinators are less mobile, decreased with increasing spatial isolation, but not that of the bumblebee-pollinated Polygonatum multiflorum. Conclusions Our study indicates that habitat loss and fragmentation compromise the long-term viability of slow-colonizing forest herbs despite their ability to persist for many decades by clonal propagation. The distinct responses of the three species studied within the same landscapes confirm the need of multi-species approaches. The mobility of associated pollinators should be considered an important determinant of forest herbs’ sensitivity to habitat loss and fragmentation.

[Context] Functional connectivity is vital for plant species dispersal, but little is known about how habitat loss and the presence of green infrastructure interact to affect both functional and structural connectivity, and the impacts of each on species groups., [Objectives] We investigate how changes in the spatial configuration of species-rich grasslands and related green infrastructure such as road verges, hedgerows and forest borders in three European countries have influenced landscape connectivity, and the effects on grassland plant biodiversity., [Methods] We mapped past and present land use for 36 landscapes in Belgium, Germany and Sweden, to estimate connectivity based on simple habitat spatial configuration (structural connectivity) and accounting for effective dispersal and establishment (functional connectivity) around focal grasslands. We used the resulting measures of landscape change to interpret patterns in plant communities., [Results] Increased presence of landscape connecting elements could not compensate for large scale losses of grassland area resulting in substantial declines in structural and functional connectivity. Generalist species were negatively affected by connectivity, and responded most strongly to structural connectivity, while functional connectivity determined the occurrence of grassland specialists in focal grasslands. Restored patches had more generalist species, and a lower density of grassland specialist species than ancient patches., [Conclusions] Protecting both species rich grasslands and dispersal pathways within landscapes is essential for maintaining grassland biodiversity. Our results show that increases in green infrastructure have not been sufficient to offset loss of semi-natural habitat, and that landscape links must be functionally effective in order to contribute to grassland diversity., This research was funded through the 2015–2016 BiodivERsA COFUND call for research proposals, with the national funders the Swedish Research Council for Environment, Agricultural Sciences and Spatial Planning (FORMAS), the Swedish Environmental Protection Agency (Naturvårdsverket), the Belgian Science Policy Office (BelSPo), the Germany Federal Ministry of Education and Research (Bundesministerium fuer Bildung und Forschung, FKZ: 01LC1619A) and the Spanish Ministry of Science, Innovation and Universities (Ministerio de Ciencia, Innovación y Universidades). Open access funding provided by Stockholm University.

Linear landscape elements such as hedgerows and road verges have the potential to mitigate the adverse effects of habitat fragmentation and climate change on species, for instance, by serving as a refuge habitat or by improving functional connectivity across the landscape. However, so far this hypothesis has not been evaluated at large spatial scales, preventing us from making generalized conclusions about their efficacy and implementation in conservation policies. Here, we assessed plant diversity patterns in 336 vegetation plots distributed along hedgerows and road verges, spanning a macro‐environmental gradient across temperate Europe. We compared herb‐layer species richness and composition in these linear elements with the respective seed‐source (core) habitats, that is, semi‐natural forests and grasslands. Next, we assessed how these differences related to several environmental drivers acting either locally, at the landscape level or along the studied macro‐ecological gradient. Across all regions, about 55% of the plant species were shared between forests and hedgerows, and 52% between grasslands and road verges. Habitat‐specialist richness was 11% lower in the linear habitats than in the core habitats, while generalist richness was 14% higher. The difference in floristic composition between both habitat types was mainly due to species turnover, and not nestedness. Most notably, forest‐specialist richness in hedgerows responded positively to tree cover, tree height and the proportion of forests in the surrounding landscape, while generalist richness was negatively affected by tree height and buffering effect of trees on subcanopy temperatures. Grassland and road verge diversity was mainly influenced by soil properties, with positive effects of basic cation levels on the number of specialists and those of bioavailable soil phosphorus on generalist diversity. Synthesis and applications. We demonstrate that linear landscape elements provide a potential habitat for plant species across Europe, including slow‐colonizing specialists. Additionally, our results stress the possibility for land managers to modify local habitat features (e.g. canopy structure, subcanopy microclimate, soil properties, mowing regime) through management practices to enhance the colonization success of specialists in these linear habitats. These findings underpin the management needed to better conserving the biodiversity of agricultural landscapes across broad geographical scales. ispartof: Journal Of Applied Ecology vol:57 issue:7 pages:1244-1257 status: Published online

Aim Available theories suggest that the species number increases with island size, but the relationship between island size and species density, the number of species per unit area, is less underst ...

Questions: Rates of plant community shifts after environmental changes depend on how quickly affected species are gained and lost. Understanding how the balance between extinction and colonisation varies over time, and how it is influenced by local and landscape factors, is essential to understanding overall change trajectories. Investigating change requires data at several time steps over sufficient periods, and the paucity of such data represents an important knowledge gap. We ask: (a) how variation over time in the rates of species’ extinction and species’ colonization controls the trajectory of biodiversity change in abandoned semi‐natural grasslands? and (b) can landscape composition and habitat history modify change trajectories by acting independently on groups within plant communities? Location: Sweden, Stockholm Archipelago. Methods: We use data on plant composition, management history and landscape context in former grasslands, abandoned at different points since 1901, in a space‐for‐time analysis, comparing rates of grassland species loss and forest species establishment and investigating resulting biodiversity trajectories. Results: Grassland species declined steeply in recently abandoned habitats before levelling off, while the accumulation of forest species was linear, with no plateau reached even at the longest time since abandonment. Hence, we observed a trough in biodiversity, with an initial decline in overall species richness followed by a partial recovery. Only forest species gain was enhanced by nearby habitat availability. Conclusions: Information on community compositional changes over short time periods may be misleading about the extent and even direction of ongoing biodiversity gains and losses. Moreover, the non‐linear changes observed suggest thresholds in time, after which succession to the forest community accelerates and the ability to manage a return to the grassland community diminishes. Accounting for the combined influence of landscape composition and history is key to fully understanding community shifts over time.

QUESTIONS: We aim to answer three questions: (a) what are the effects of canopy shading of different tree species on seed germination of eight understorey grassland species? (b) how is plant species’ occurrence in understorey communities affected by differences in canopy cover and does that depend on tree species composition? and (c) are there mechanistic links between the effects of trees on species’ germination and their occurrence in the understorey plant community? LOCATION: Semi‐natural wooded pastures in the biosphere reserve Ostra Vatterbranterna, southern Sweden. METHODS: In this study, we examined the germination of eight grassland plant species in a seed sowing experiment under natural conditions in the field. Seeds were sown beneath and outside the canopy of two tree species within 48 plots split up by four wooded pasture sites. We combined observed germination responses with a plant community survey to assess the effects of canopy cover in relation to tree species composition on plant community responses. We analysed these data in relation to species’ seed mass and vegetative shade tolerance. RESULTS: Shade‐tolerant species germinated better beneath compared to outside tree canopies, without any clear advantage of large‐seeded over small‐seeded species. As expected, species’ shade tolerance was also positively related to canopy cover within the understorey plant community. Importantly, we found strong tree species‐specific effects of canopy shading on the species’ germination response, but not on their presence within the plant community. However, optimal canopy cover conditions for germination and for the mature plants differed across grassland species and depended on tree species. CONCLUSIONS: Our results show that different tree species play key ecosystem engineering roles in shaping wooded grassland plant community composition at the germination stage. Management practices favouring specific tree species may therefore be highly relevant for targeted biodiversity conservation of wooded semi‐natural grasslands.

© 2019 The Authors. Diversity and Distributions Published by John Wiley & Sons Ltd Aim: Forests are highly fragmented across Western Europe, making forest edges important features in many agricultural landscapes. Forest edges are subject to strong abiotic gradients altering the forest environment and resulting in strong biotic gradients. This has the potential to change the forest's capacity to provide multiple ecosystem services such as nutrient cycling, carbon sequestration and natural pest control. Soil organisms play a key role in this perspective; however, these taxa are rarely considered in forest edge research. Location: A latitudinal gradient of 2,000 km across Western Europe. Methods: We sampled six dominant taxa of litter-dwelling macro-arthropods (carabid beetles, spiders, harvestmen, centipedes, millipedes and woodlice) in forest edges and interiors of 192 forest fragments in 12 agricultural landscapes. We related their abundance and community composition to distance from the edge and the interaction with forest age, edge orientation and edge contrast (contrast between land use types at either side of the edge). Results: Three out of six macro-arthropod taxa have higher activity-density in forest edges compared to forest interiors. The abundance patterns along forest edge-to-interior gradients interacted with forest age. Forest age and edge orientation also influenced within-fragment compositional variation along the forest edge-to-interior gradient. Edge contrast influenced abundance gradients of generalist predators. In general, older forest fragments, south-oriented edges and edges along structurally more continuous land use (lower contrast between forest and adjacent land use) resulted in stronger edge-to-interior gradients while recent forests, north-oriented edges and sharp land use edges induced similarity between forest edge and interior along the forest edge-to-interior gradients in terms of species activity-density and composition. Main conclusions: Edge effects on litter-dwelling macro-arthropods are anticipated to feedback on important ecosystem services such as nutrient cycling, carbon sequestration and natural pest control from small forest fragments. ispartof: DIVERSITY AND DISTRIBUTIONS vol:25 issue:6 pages:963-974 status: published

Semi-natural grasslands harbour many of Europe's species of conservation interest. Although larger grasslands are the focus of most conservation activity, many grassland fragments are scattered across landscapes –in small patches or along linear elements– which can form Grassland Green Infrastructure (GGI). GGI has the potential to enhance landscape diversity by creating functioning metacommunities comprising of large semi-natural grasslands and these surrounding fragments. While often highlighted in conservation policy, little is known about the biodiversity supported by green infrastructure itself and thus its conservation potential. To address this issue, we contrasted plant communities in 36 ‘core’ grassland sites across three European countries with communities in the surrounding GGI. We related compositional differences to amount and type of GGI habitat (patches or linear), and the distances for seed dispersal by livestock from core sites. We found substantial differences between the GGI and the core sites, with a mean 54% species turn-over. These differences indicated filtering of stress tolerant species characteristic of low nutrient conditions, and semi-natural grassland specialists. Species with poorer dispersal abilities declined strongly with increasing distances from the core sites. The many additional species in the GGI, not found in the core sites, were predominantly those with a competitive strategy and high seed dispersal ability. We conclude that the biodiversity-supporting role of GGI across Europe is severely constrained by eutrophication, dispersal limitation and external propagule pressure. Actions to improve the quality of GGI might include enhancing dispersal by livestock combined with more type-diversification and less intensively used grassland habitats., This research was funded through the 2015-2016 BiodivERsA COFUND call for research proposals, with the national funders FORMAS (2016-01948), the Swedish Environmental Protection Agency (Naturvårdsverket), the Belgian Science Policy Office (BelSPo, BR/175/A1/FUNgreen), the Germany Federal Ministry of Education and Research (Bundesministerium fuer Bildung und Forschung, FKZ: 01LC1619A) and the Spanish Ministry of Science, Innovation and Universities (Ministerio de Ciencia, Innovación y Universidades, PCIN-2016-077). JMB was funded under UKCEH National Capability project 06895.

International audience; Forest biodiversity world-wide is affected by climate change, habitat loss and fragmentation, and today 20% of the forest area is located within 100 m of a forest edge. Still, forest edges harbour a substantial amount of terrestrial biodiversity, especially in the understorey. The functional and phylogenetic diversity of forest edges have never been studied simultaneously at a continental scale, in spite of their importance for the forests' functioning and for communities' resilience to future change. We assessed nine metrics of taxonomic, phylogenetic and functional diversity of understorey plant communities in 225 plots spread along edge-to-interior gradients in deciduous forests across Europe. We then derived the relative effects and importance of edaphic, stand and landscape conditions on the diversity metrics. Here, we show that taxonomic, phylogenetic and functional diversity metrics respond differently to environmental conditions. We report an increase in functional diversity in plots with stronger microclimatic buffering, in spite of their lower taxonomic species richness. Additionally, we found increased taxonomic species richness at the forest edge, but in forests with intermediate and high openness, these communities had decreased phylogenetic diversity. Functional and phylogenetic diversity revealed complementary and important insights in community assembly mechanisms. Several environmental filters were identified as potential drivers of the patterns, such as a colder macroclimate and less buffered microclimate for functional diversity. For phylogenetic diversity, edaphic conditions were more important. Interestingly, plots with lower soil pH had decreased taxonomic species richness, but led to increased phylogenetic diversity, challenging the phylogenetic niche conservatism concept. Synthesis. Taxonomic, phylogenetic and functional diversity of understorey communities in forest edges respond differently to environmental conditions, providing insight into different community assembly mechanisms and their interactions. Therefore, it is important to look beyond species richness with phylogenetic and functional diversity approaches when focusing on forest understorey biodiversity.

Global environmental changes are expected to alter the functional characteristics of understorey herb-layer communities, potentially affecting forest ecosystem functioning. However, little is known about what drives the variability of functional traits in forest understories. Here, we assessed the role of different environmental drivers in shaping the functional trait distribution of understorey herbs in fragmented forests across three spatial scales. We focused on 708 small, deciduous forest patches located in 16 agricultural landscape windows, spanning a 2500-km macroclimatic gradient across the temperate forest biome in Europe. We estimated the relative effect of patch-scale, landscape-scale and macroclimatic variables on the community mean and variation of plant height, specific leaf area and seed mass. Macroclimatic variables (monthly temperature and precipitation extremes) explained the largest proportion of variation in community trait means (on average 77% of the explained variation). In contrast, patch-scale factors dominated in explaining community trait variation (on average 68% of the explained variation). Notably, patch age, size and internal heterogeneity had a positive effect on the community-level variability. Landscape-scale variables explained only a minor part of the variation in both trait distribution properties. The variation explained by shared combinations of the variable groups was generally negligible. These findings highlight the importance of considering multiple spatial scales in predictions of environmental-change effects on the functionality of forest understories. We propose that forest management sustainability could benefit from conserving larger, historically continuous and internally heterogeneous forest patches to maximise ecosystem service diversity in rural landscapes. (C) 2018 Gesellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.

Aim Habitat loss threatens plant diversity globally. Lack of plant functional connectivity between isolated populations is often pinpointed as one of the major underlying mechanisms driving subsequ ...

Small semi-natural and natural habitats in agricultural landscapes are important for biodiversity. With modern and more intensive agricultural practices they have become smaller (less than 1600 m2) ...

Metacommunity theory emphasizes that seed dispersal not only limits but equally maintains plant diversity, though the latter receives little empirical attention. Discerning the temporal and spatial ...

Background and Aims Habitat fragmentation threatens global biodiversity. Many plant species persist in habitat fragments via persistent life cycle stages such as seed banks, generating a species extinction debt. Here, seed banks are hypothesized to cause a temporal delay in the expected loss of genetic variation, which can be referred to as a genetic extinction debt, as a possible mechanism behind species extinction debts. Methods Fragmented grassland populations of Campanula rotundifolia were examined for evidence of a genetic extinction debt, investigating if the seed bank contributed to the extinction debt build-up. The genetic make-up of 15 above- and below-ground populations was analysed in relation to historical and current levels of habitat fragmentation, both separately and combined. Key Results Genetic diversity was highest in above-ground populations, though below-ground populations contained 8 % of unique alleles that were absent above-ground. Above-ground genetic diversity and composition were related to historical patch size and connectivity, but not current patch characteristics, suggesting the presence of a genetic extinction debt in the above-ground populations. No such relationships were found for the below-ground populations. Genetic diversity measures still showed a response to historical but not present landscape characteristics when combining genetic diversity of the above- and below-ground populations. Conclusions The fragmented C. rotundifolia populations exhibited a genetic extinction debt. However, the role of the seed banks in the build-up of this extinction debt is probably small, since the limited, unique genetic diversity of the seed bank alone seems unable to counter the detrimental effects of habitat fragmentation on the population genetic structure of C. rotundifolia.

ContextThe habitat amount hypothesis has rarely been tested on plant communities. It remains unclear how habitat amount affect species richness in habitat fragments compared to island effects such ...

Understanding the responses of invasive and native populations to environmental change is crucial for reliable predictions of invasions in the face of global change. While comparisons of responses across invasive species with different life histories have been performed before, comparing functional traits of congeneric native and invasive species may help to reveal driving factors associated with invasion. Here we compared morphological functional trait patterns of an invasive species (Impatiens parviflora) with its congeneric native species (I. noli-tangere) along an approximately 1600 km European latitudinal gradient from France (49°34′N) to Norway (63°40′N). Soil nitrogen was recorded during six weeks of the growing season, and light, soil moisture, and nutrient availability were estimated for each sampled population using community weighted means of indicator values for co-occurring species. Temperature data were gathered from nearby weather stations. Both the native and invasive species are taller at higher latitudes and this response is strongest in the invasive species. Seed mass and number of seeds per capsule increase in I. noli-tangere but decrease in I. parviflora towards higher latitudes. Surprisingly, plant height in the invasive I. parviflora decreases with increasing soil nitrogen availability. The latitudinal pattern in seed mass is positively related to temperature in I. noli-tangere and negatively in I. parviflora. Leaf area of both species decreases with increasing Ellenberg indicator values for nitrogen and light but increases with increasing soil moisture. Soil nitrogen concentrations and Ellenberg indicator values for nitrogen have significant positive (I. noli-tangere) and negative (I. parviflora) effects on the number of seeds per capsule. Our results show that the native I. noli-tangere has efficient reproduction at its range edge while the invasive I. parviflora shows a marked decrease in seed size and seed number per capsule. These patterns are unrelated to the growth and obtained size of the plants: even low soil nitrogen availability in the north seemed not to limit plant growth and size. Our results suggest that the invasive I. parviflora tends to become more invasive at lower latitudes by producing heavier seeds and more seeds per capsule.

Changes in abiotic factors along altitudinal and latitudinal gradients cause powerful environmental gradients. The topography of alpine areas generates environmental gradients over short distances, and alpine areas are expected to experience greater temperature increase compared to the global average. In this study, we investigate alpha, beta, and gamma diversity, as well as community structure, of vascular plant communities along altitudinal gradients at three latitudes in the Swedish mountains. Species richness and evenness decreased with altitude, but the patterns within the altitudinal gradient varied between sites, including a sudden decrease at high altitude, a monotonic decrease, and a unimodal pattern. However, we did not observe a decline in beta diversity with altitude at all sites, and plant communities at all sites were spatially nested according to some other factors than altitude, such as the availability of water or microtopographic position. Moreover, the observed diversity patterns did not follow the latitudinal gradient. We observed a spatial modularity according to altitude, which was consistent across sites. Our results suggest strong influences of site‐specific factors on plant community composition and that such factors partly may override effects from altitudinal and latitudinal environmental variation. Spatial variation of the observed vascular plant communities appears to have been caused by a combination of processes at multiple spatial scales., We studied the altitude variation of alpine plant communities in the Swedish mountains. We found that species richness and evenness decreased with altitude but the patterns within the altitude gradient varied between sites, including a sudden decrease at high altitude, a monotonic decrease, and a unimodal pattern.

Background and Aims It remains unclear whether invasive species can maintain both high biomass and reproductive output across their invaded range. Along latitudinal gradients, allocation theory predicts that faster flowering onset at high latitudes results in maturation at smaller size and thus reduced reproductive output. For annual invasive plants, more favourable environmental conditions at low latitudes probably result in stronger competition of co-occurring species, potentially driving selection for higher investment in vegetative biomass, while harsher climatic conditions and associated reproductive uncertainty at higher latitudes could reduce selection for vegetative biomass and increased selection for high reproductive investment (stress-gradient hypothesis). Combined, these drivers could result in increased or constant reproductive allocation with increasing latitude. Methods We quantified life-history traits in the invasive annual plant Impatiens glandulifera along a latitudinal gradient in Europe. By growing two successive glasshouse generations, we assessed genetic differentiation in vegetative growth and reproductive output across six populations, and tested whether onset of flowering drives this divergence. Key Results Trait variation was mainly caused by genetic differentiation. As expected, flowering onset was progressively earlier in populations from higher latitudes. Plant height and vegetative biomass also decreased in populations from higher latitudes, as predicted by allocation theory, but their variation was independent of the variation in flowering onset. Reproductive output remained constant across latitudes, resulting in increased reproductive allocation towards higher latitudes, supporting the stress-gradient hypothesis. We also observed trait genetic differentiation among populations that was independent of latitude. Conclusions We show that an annual invasive plant evolved several life-history traits across its invaded range in ~150 years. The evolution of vegetative and reproductive traits seems unconstrained by evolution of flowering onset. This genetic decoupling between vegetative and reproductive traits possibly contributes to the invasion success of this species.

Question Belowground processes are key determinants of aboveground plant population and community dynamics. Still, our understanding of how environmental drivers shape plant communities is mostly based on aboveground diversity patterns, bypassing belowground plant diversity stored in seed banks. As seed banks may shape aboveground plant communities, we question whether concurrently analyzing the above- and belowground species assemblages may potentially enhance our understanding of community responses to environmental variation. Location Temperate deciduous forests along a 2000 km latitudinal gradient in NW-Europe. Methods Herb layer, seed bank and local environmental data including soil pH, canopy cover, forest cover continuity and time since last canopy disturbance were collected in 129 temperate deciduous forest plots. We quantified herb layer and seed bank diversity per plot and evaluated how environmental variation structured community diversity in the herb layer, seed bank and the combined herb layer-seed bank community. Results Seed banks consistently held more plant species than the herb layer. How local plot diversity was partitioned across the herb layer and seed bank was mediated by environmental variation in drivers serving as proxies of light availability. The herb layer and seed bank contained an ever smaller and ever larger share of local diversity, respectively, as both canopy cover and time since last canopy disturbance decreased. Species richness and β-diversity of the combined herb layer-seed bank community responded distinctly different compared to the separate assemblages in response to environmental variation in, e.g., forest cover continuity and canopy cover. Conclusions The seed bank is a belowground diversity reservoir of the herbaceous forest community, which interacts with the herb layer, though constrained by environmental variation in e.g. light availability. The herb layer and seed bank coexist as a single community by means of the so-called storage effect, resulting in distinct responses to environmental variation not necessarily recorded in the individual herb layer or seed bank assemblages. Thus, concurrently analysing above- and belowground diversity will improve our ecological understanding of how understorey plant communities respond to environmental variation. This article is protected by copyright. All rights reserved.

African elephants are ‘keystone’ species with respect to biodiversity conservation in Africa since they maintain habitats that support several animal communities by changing vegetation structure through foraging and by dispersing seeds between landscapes. Elephants are also ‘flagship’ species because, given their impressive size, they can make people sympathetic and stimulate local and international concerns for their protection. Economically, elephants contribute to national revenues as tourists are willing to pay to watch them. Despite all these factors, little is known however about elephant movement and how they utilize resources, especially in miombo-wetland ecosystems. This thesis investigates how elephants utilize resources in a miombo-wetland ecosystem in the Ugalla landscape of Western Tanzania over different protected areas containing different resource users. Using Global Positioning System (GPS) collars fitted to six elephants, it was observed that some elephant families are not confined in one protected area in the Ugalla landscape. Rather, they moved readily between different protected areas. Elephant movements were restricted to areas near the rivers, especially the Ugalla River, during the dry season and were dispersed widely during the wet season. As they move, elephants in the miombo woodlands of Ugalla selected the most abundant woody plants for browsing. Common to many woody plants, the browsed plants were short of mineral nutrients (e.g., sodium, calcium). Elephants obtained additional minerals by eating soils from certain termite mounds. Soils from termite mounds are richer in mineral elements (e.g., sodium, calcium, iron) compared to soils from the surrounding flood plain or compared to the browsed plants. However, the recorded termite mounds from which elephants eat soils were not evenly distributed in the landscape but confined mainly to the flood plains in the Ugalla Game Reserve. The Ugalla River, which is the main source of water for the elephants and other animals and also supports fishing activities by the local people in Ugalla during the dry seasons, is infested by the water hyacinth (Eichhornia crassipes). Such infestation potentially limits access to these precious surface water supplies. In addition at the regional level, the Ugalla River is among the major rivers that flow into the Lake Tanganyika which is shared by the countries of Tanzania, Burundi, Democratic Republic of Congo and Zambia. Thus, the spread of water hyacinth if left unchecked threatens to impact Lake Tanganyika, affecting many countries and ecosystem services. This thesis highlights that sustainable conservation of biodiversity in different protected areas in the Ugalla landscape requires an integrated management approach that will embrace conservation of different interrelated landscape resources required by both wildlife and the rural poor populations for their livelihoods. Regular coordinated wildlife anti-poaching patrols should be initiated across the entire Ugalla landscape because the elephants, among other wildlife, utilize different protected areas in Ugalla. Local communities should also be engaged in conservation initiatives (e.g., controlling the spread of the water hyacinth) as these directly impact local livelihoods.

The soil microbial community is essential for maintaining ecosystem functioning and is intimately linked with the plant community. Yet, little is known on how soil microbial communities in the root zone vary at continental scales within plant species. Here we assess the effects of soil chemistry, large-scale environmental conditions (i.e. temperature, precipitation and nitrogen deposition) and forest land-use history on the soil microbial communities (measured by phospholipid fatty acids) in the root zone of four plant species (Geum urbanum, Milium effusum, Poa nemoralis and Stachys sylvatica) in forests along a 1700 km latitudinal gradient in Europe. Soil microbial communities differed significantly among plant species, and soil chemistry was the main determinant of the microbial community composition within each plant species. Influential soil chemical variables for microbial communities were plant species-specific; soil acidity, however, was often an important factor. Large-scale environmental conditions, together with soil chemistry, only explained the microbial community composition in M. effusum and P. nemoralis. Forest land-use history did not affect the soil microbial community composition. Our results underpin the dominant role of soil chemistry in shaping microbial community composition variation within plant species at the continental scale, and provide insights into the composition and functionality of soil microbial communities in forest ecosystems.

Phenotypic trait variation plays a major role in the response of plants to global environmental change, particularly in species with low migration capabilities and recruitment success. However, little is known about the variation of functional traits within populations and about differences in this variation on larger spatial scales. In a first approach, we therefore related trait expression to climate and local environmental conditions, studying two temperate forest herbs, Milium effusum and Stachys sylvatica, along a similar to 1800-2500 km latitudinal gradient. Within each of 9-10 regions in six European countries, we collected data from six populations of each species and recorded several variables in each region (temperature, precipitation) and population (light availability, soil parameters). For each plant, we measured height, leaf area, specific leaf area, seed mass and the number of seeds and examined environmental effects on within-population trait variation as well as on trait means. Most importantly, trait variation differed both between and within populations. Species, however, differed in their response. Intrapopulation variation in Milium was consistently positively affected by higher mean temperatures and precipitation as well as by more fertile local soil conditions, suggesting that more productive conditions may select for larger phenotypic variation. In Stachys, particularly light availability positively influenced trait variation, whereas local soil conditions had no consistent effects. Generally, our study emphasises that intra-population variation may differ considerably across larger scales-due to phenotypic plasticity and/or underlying genetic diversity-possibly affecting species response to global environmental change.

Questions How does small linear grassland elements (paddy levees and road verges) contribute to plant diversity patterns in sites with modern or traditional management? Which factors determine small-scale plant species richness of small linear grassland elements in a Japanese agricultural landscape? Location Shiga Prefecture, Japan. Methods Aerial photos from 1963, 1985, 1995 and 2008 were used to investigate landscape changes and to measure landscape structures within a typical agricultural landscape (~25 km2). Plant diversity was investigated in ten paddy levees and ten road verges in each of four traditional (barely any land cover alterations after 1963) and five modern (almost completely changed after 1985) sites. α-diversity (mean number of species per plot) and γ-diversity (total number of species) for all species, specialist and invasive species, respectively, were analysed in relation to site history types and landscape structures. Plot species richness was analysed in relation to site history type, patch age and width, proximity to forest and litter removal. Results In total, 204 plant species were found, 55 species unique to traditional sites and 13 unique to modern sites. α- and γ-diversity of paddy levees and road verges was significantly higher in traditional sites. Number of specialist species on paddy levees and number of invasive species in road verges was also higher in traditional sites. Age was the strongest predictor of plot species richness on levees; older levees supported higher species richness than younger ones. Litter removal was the strongest predictor (positive effect) of plot species richness in road verges. Conclusions These results suggest that the colonization credit of new levees in modern sites is decreasing quite rapidly as species quickly establish, approaching almost the same richness values as in traditional sites. However, long continuity of traditional management is still one of the most important contributors to plant diversity and crucial for many species. But small grassland elements can still be of great significance for preserving and enhancing plant diversity in otherwise depleted landscapes, as long as they are managed and connected throughout the landscape.

Aim Macroclimate is a major determinant of large-scale diversity patterns. However, the influence of smaller-scale factors on local diversity across large spatial extents is not well documented. Here, we quantify the relative importance of local (patch-scale), landscape-scale and macroclimatic drivers of herbaceous species diversity in small forest patches in agricultural landscapes across Europe. Location Deciduous forest patches in eight regions along a macroclimatic gradient from southern France to central Sweden and Estonia. Methods The diversity of forest specialists and generalists at three levels (whole forest patch, sampling plots within patches and between scales) was related to patch-scale (forest area, age, abiotic and biotic heterogeneity), landscape-scale (amount of forest, grasslands and hedgerows around the patch, patch isolation) and macroclimatic variables (temperature and precipitation) using generalized linear mixed models and variation partitioning for each group of variables. Results The total amount of explained variation in diversity ranged from 8% for plot-scale diversity of generalists to 54% for patch-scale diversity of forest specialists. Patch-scale variables always explained more than 60% of the explained variation in diversity, mainly due to the positive effect of within-patch heterogeneity on patch-scale and between-scale diversities and to the positive effect of patch age on plot-scale diversity of forest specialists. Landscape-scale variables mainly contributed to the amount of explained variation in plot-scale diversity, being more important for forest specialists (21%) than for generalists (18%). Macroclimatic variables contributed a maximum of 11% to the plot-scale diversity of generalists. Main conclusions Macroclimate poorly predicts local diversity across Europe, and herbaceous diversity is mainly explained by habitat features, less so by landscape structure. We show the importance of conserving old forest patches as refugia for typical forest species, and of enhancing the landscape context around the patches by reducing the degree of disturbance caused by agriculture.

The rare forest plant Chimaphila umbellata (Ericaceae) has decreased drastically during the last century, approximately by 80 % in some regions in Sweden. We examined associations between various biotic and abiotic conditions related to changes in forest management and nitrogen deposition, and C. umbellata population size, flowering frequency, fruit set and seed production. Environmental conditions at 38 C. umbellata sites in the provinces of Uppland and Sodermanland, Sweden, included light inflow, cover of competitive species, soil nitrogen, continuity of forest cover and soil texture. The results suggested that population size was negatively affected by cover of competitive species. Population size was not related to light availability although increased shading was associated with decreased flowering frequency. Fruit set was negatively affected by cover of competitive species, and seed production decreased with increasing soil nitrogen content. Fruit set and seed production increased with increasing population size. This study shows that denser forest stands and increased abundance of Vaccinium myrtillus and graminoid species may have strong negative effects on C. umbellata. This species’ longevity and clonal propagation may buffer some negative impacts, which in turn might contribute to an extinction debt. To maintain viable populations of Chimaphila umbellata in Swedish forests, there is a need for targeted management in forest habitats, i.e. reintroducing moderate disturbance regimes to reduce competition and increase light inflow.

Semi-natural grasslands have been severely affected by agricultural land-use change. However, the isostatic land adjustment following deglaciation in the Northern Hemisphere means that new land is continually being created in coastal areas. We modelled isostatic adjustment during the last 4000 years in a region of the Baltic coast to estimate the emergence of potential grassland habitat. We also compared the α and β diversity of existing managed and abandoned coastal meadows, and assessed their contribution to biodiversity at landscape scales. We estimated that half the 7866 km 2 of emerging land had the potential to become coastal meadow habitat, which is an order of magnitude larger than the total area of all valuable semi-natural grassland in the study region today. The small area of managed coastal habitat remaining was found to have a disproportionate influence on the richness of threatened species at landscape scales, but our results also show that continued management is essential for the maintenance of grassland biodiversity. Our combination of approaches identifies uplifted coastal meadows as an additional original niche for grassland plant species, while highlighting that low-intensity disturbance through grassland management is essential for the maintenance of diversity at multiple scales.

Dispersal is a key process in determining the survival of plant species following habitat fragmentation and climate change, as well as driving the introduction and spread of invasive alien species in new regions. Due to its passive nature, seed dispersal is particularly complex, and the rare long-distance events relevant for plant species' responses to environmental change are a barrier to its understanding. Attempts to simplify the seed dispersal process often ignore dispersal by humans, despite the huge influence humans have over ecological systems throughout the world. In this Biodiversity Viewpoint, we describe how the movement patterns of humans and human-mediated dispersal vectors can be useful for understanding potential patterns of dispersal at multiple spatial scales. Humans and their associated dispersal vectors such as livestock and motor vehicles can disperse huge numbers of seeds of many plant species very long distances. Their relationships with the physical environment affect their movement, and therefore the movement of the seeds which they can potentially disperse. Therefore, we believe that a geographical approach can be useful at a time where understanding and managing pathways of dispersal are of direct relevance to the challenges faced by plant species and communities.

Climate change will likely affect population dynamics of numerous plant species by modifying several aspects of the life cycle. Because plant regeneration from seeds may be particularly vulnerable, here we assess the possible effects of climate change on seed characteristics and present an integrated analysis of seven seed traits (nutrient concentrations, samara mass, seed mass, wing length, seed viability, germination percentage, and seedling biomass) of Acer platanoides and A. pseudoplatanus seeds collected along a wide latitudinal gradient from Italy to Norway. Seed traits were analyzed in relation to the environmental conditions experienced by the mother trees along the latitudinal gradient. We found that seed traits of A. platanoides were more influenced by the climatic conditions than those of A. pseudoplatanus. Additionally, seed viability, germination percentage, and seedling biomass of A. platanoides were strongly related to the seed mass and nutrient concentration. While A. platanoides seeds were more influenced by the environmental conditions (generally negatively affected by rising temperatures), compared to A. pseudoplatanus, A. platanoides still showed higher germination percentage and seedling biomass than A. pseudoplatanus. Thus, further research on subsequent life-history stages of both species is needed. The variation in seed quality observed along the climatic gradient highlights the importance of studying the possible impact of climate change on seed production and species demography.

Global warming and enhanced nitrogen (N) inputs are two key global-change drivers affecting temperate forest ecosystems simultaneously. Interactive effects of multiple drivers might cause species responses to differ from those in single-factor experiments; therefore, there is an urgent need for more multi-factor studies. Here, we assessed the growth and reproductive performance of multiple populations of a widespread grass of deciduous forests (Milium effusum) sampled along a latitudinal gradient and subjected to experimental manipulations of temperature and nitrogen availability. Common garden transplant experiments along the latitudinal gradient were used to manipulate temperatures and combined with experimental N addition to assess intraspecific responses of the study species to global-change drivers as well as to determine local adaptation. The total biomass, number of seeds and seedling emergence time of M. effusum increased when transplanted in the southern common garden. Apart from effects on the seed mass, the species did not respond to N addition alone. Yet, interactive effects between warming and N addition were found: N addition led to increased biomass growth but only in the northern common garden. Significant home-site advantages were apparent, most likely because of increased mycorrhizal colonization of roots of local transplants. We show that multiple global-change drivers may alter dynamics in understorey communities of temperate forests. Our study reinforces the need to increase our understanding of plant responses to future environmental changes by expanding the multi-factor research framework.

Climate change is acting on several aspects of plant life cycles, including the sexual reproductive stage, which is considered amongst the most sensitive life-cycle phases. In temperate forests, it is expected that climate change will lead to a compositional change in community structure due to changes in the dominance of currently more abundant forest tree species. Increasing our understanding of the effects of climate change on currently secondary tree species recruitment is therefore important to better understand and forecast population and community dynamics in forests. Here, we analyse the interactive effects of rising temperatures and soil moisture reduction on germination, seedling survival and early growth of two important secondary European tree species, Acer pseudoplatanus and A. platanoides. Additionally, we analyse the effect of the temperature experienced by the mother tree during seed production by collecting seeds of both species along a 2200-km long latitudinal gradient. For most of the responses, A. platanoides showed higher sensitivity to the treatments applied, and especially to its joint manipulation, which for some variables resulted in additive effects while for others only partial compensation. In both species, germination and survival decreased with rising temperatures and/or soil moisture reduction while early growth decreased with declining soil moisture content. We conclude that although A. platanoides germination and survival were more affected after the applied treatments, its initial higher germination and larger seedlings might allow this species to be relatively more successful than A. pseudoplatanus in the face of climate change.

The extensive transformation of agricultural landscapes worldwide has led to a decrease in grassland species related to traditional low-intensive farming. To properly manage and protect species, habitats and ecosystems in any of these landscapes requires a better understanding of direct and indirect effects of the processes driving biodiversity decline. In this study, we investigated how small habitat elements, represented by mid-field islets and road verges, in different types of agricultural landscapes can sustain a regional species pool for plant diversity otherwise associated to semi-natural grasslands. Although semi-natural grasslands had higher total and specialist richness, we found that small habitat elements harboured relatively high plant species richness, especially in a landscape with many semi-natural grasslands left. In the most intensively managed landscape, total richness declined as distance to the nearest semi-natural grassland increased. In contrast, β-diversity was highest in these landscapes indicating that small habitats are also negatively affected by distance to grassland. We found that species trait composition varied depending on habitat and landscape. The results confirm that fragmentation shape trait composition within plant communities, e.g. plant size, clonality, longevity, and dispersal traits. We conclude that small habitat elements increase the total area available to grassland species present in the landscape, boosting the spatio-temporal dynamics of grassland communities. Small habitat elements may hence function as refugia or stepping stone habitats, especially in intensively utilized agricultural landscapes, and should be regarded as a functional part of a semi-natural grassland network, analogous to a meta-population.

Aim In response to environmental changes and to avoid extinction, species may either track suitable environmental conditions or adapt to the modified environment. However, whether and how species adapt to environmental changes remains unclear. By focusing on the realized niche (i.e. the actual space that a species inhabits and the resources it can access as a result of limiting biotic factors present in its habitat), we here examine shifts in the realized-niche width (i.e. ecological amplitude) and position (i.e. ecological optimum) of 26 common and widespread forest understorey plants across their distributional ranges. Location Temperate forests along a ca. 1800-km-long latitudinal gradient from northern France to central Sweden and Estonia. Methods We derived species' realized-niche width from a β-diversity metric, which increases if the focal species co-occurs with more species. Based on the concept that species' scores in a detrended correspondence analysis (DCA) represent the locations of their realized-niche positions, we developed a novel approach to run species-specific DCAs allowing the focal species to shift its realized-niche position along the studied latitudinal gradient while the realized-niche positions of other species were held constant. Results None of the 26 species maintained both their realized-niche width and position along the latitudinal gradient. Few species (9 of 26: 35%) shifted their realized-niche width, but all shifted their realized-niche position. With increasing latitude, most species (22 of 26: 85%) shifted their realized-niche position for soil nutrients and pH towards nutrient-poorer and more acidic soils. Main conclusions Forest understorey plants shifted their realized niche along the latitudinal gradient, suggesting local adaptation and/or plasticity. This macroecological pattern casts doubt on the idea that the realized niche is stable in space and time, which is a key assumption of species distribution models used to predict the future of biodiversity, hence raising concern about predicted extinction rates.

Aim Seed banks are central to the regeneration strategy of many plant species. Any factor altering seed bank density thus affects plant regeneration and population dynamics. Although seed banks are ...

In addition to habitat loss and fragmentation, agricultural change has led to a change in seed dispersal processes in therural landscape through a loss of structural and functional connectivity. He ...

Despite a good understanding on how dispersal in space structures plant communities in fragmented landscapes, we know little about dispersal in time. Empirical evidence on temporal dispersal – the soil seed bank – is lacking, with only trait-based evidence on the seed banks’ importance for species persistence in fragmented landscapes. Therefore, seed banks of remnant grassland fragments were analyzed in how they changed compared to semi-natural grasslands following fragmentation. We studied the historical trajectories in time since fragmentation, fragment size and habitat quality of 134 grassland plots, linking these to their seed bank and plant community to understand how seed banks temporally connect grassland fragments, potentially conserving the flora of historically large semi-natural grasslands. Seed-banking grassland species were present in similar proportions in all remnant grassland fragments. The seed bank composition changed with time since fragmentation started, triggered by the deterministic loss of grassland species, generating nested subsets of the seed banks of semi-natural grasslands. The spatial heterogeneity in seed bank composition among grassland fragments limited the loss of grassland species at the landscape scale. The seed bank became an increasingly important constituent of total plant diversity with time since fragmentation started, as grassland species stored an increasingly larger proportion of their local diversity in the seed bank. Temporal dispersal enables the prolonged presence and persistence of numerous typical grassland species in fragmented landscapes. The seed banks’ storage effect of plant diversity is of considerable significance to efforts aimed at conserving and restoring plant diversity in fragmented landscapes.

Aim We investigated how current and historical land use and landscape structure affect species richness and the processes of extinction, immigration and species turnover. Location The northern part ...

Many recent studies have explored the effects of present and past landscape structure on species distribution and diversity. However, we know little about the effects of past landscape structure on ...

Summary 1. Theoretical models show that environmental heterogeneity and dispersal are major determinants of species diversity at multiple scales, yet there are few studies from real landscapes that adequately integrate variation in the surrounding matrix. Understanding how landscape context and management influence species composition and diversity patterns across habitats and scales is an important goal in ecology with relevance for both management and conservation. 2. We used a system of 25 landscapes distributed across islands in the Baltic Sea to investigate the effect of current and historical landscape context and management on plant diversity and composition in grassland communities. Plant diversity was measured at three hierarchical scales (1 m2, habitat, landscape) in grazed fields and adjacent wood pastures to calculate α-, β- and γ-diversity values across habitats and scales. 3. Structural equation modelling was used to model and quantify the effects of landscape context on species diversity and spatial turnover, and constraint analysis of principal coordinates to relate variation in species composition to landscape variables. 4. Proportion of open land, spacing and grazing intensity positively affected species diversity in both habitats, whereas the effect of historical landscape context was only significant in open fields. Plant diversity in field pastures was mainly determined by the number of species found at a small scale, while both local species density and spatial turnover were key determinants of diversity in wood pastures. 5. Habitat proximity influenced species composition as compositional similarity was higher between adjacent field and wood pastures compared to randomly paired habitats. Although increasing flow of propagules from adjacent patches can promote local coexistence, dispersal can result in spatial homogenization. 6. Synthesis. Plant diversity in grassland communities is substantially influenced by species occurring in adjacent habitats. While the effect of landscape context and management on small-scale diversity was consistent across habitats, the effect on spatial turnover was habitat specific. Our study shows that plant diversity is structured through the interplay between local and landscape processes and highlights that plant communities in specific habitat types cannot be considered in isolation from the surrounding landscape matrix.

Question To what extent does the movement of animals between fragmented habitat patches provide functional connectivity via endozoochorous seed dispersal? Location The Stockholm archipelago, Sweden. Methods We followed all movements of livestock between islands during one grazing season. After each movement, manure was collected and its seed content assessed through seedling emergence. Seedling data were then compared to vegetation surveys from the grazed islands with regard to functional traits. Results Light- and nitrogen-demanding locally abundant species, and those with relatively small and persistent seeds were more likely to be moved between islands. For quantitative traits, only a subset of the available trait ranges were dispersed, with extreme values left behind. Species apparently specialized to other means of dispersal emerged from the manure samples. Neither dispersed traits nor seed density changed with timing of movement, but seed richness and diversity both increased throughout the season. The subsets of endozoochorously-dispersed species in the established vegetation were more similar than non-dispersed subsets between islands linked by livestock. Conclusions Grazing networks contribute to the connectivity of the core species in the system, and could provide useful tools for grassland management in fragmented landscapes.

The importance of the landscape surrounding a protected area for sustaining its values is frequently discussed in conservation literature. Studies on the interactions of urbanisation and nature conservation at the global scale suggest that protected nature attracts urbanisation, and that this in turn might negatively impact the area. However, studies specifically addressing such land use dynamics at city scale are largely missing. In this study we examine the change in proportion of built up area in two zones (500 m and 1000 m) surrounding 15 urban nature reserves in southern Sweden. By using comprehensive maps from the last 50 years, we compared the zones to the overall urbanisation in the cities to reveal discrepancies in land use surrounding the nature reserves. We found that the amount of built up area in the buffer zones surrounding nature reserves followed the same trend as the corresponding cities and this relation was stable over time, although the positive relationship was not significant. The establishment of nature reserves had no detectable effect on surrounding land use, however two distinguished groups of reserves were identified with either more or less built up area in buffers zones compared to cities. These differences were related to specific local drivers such as land ownership, land use history and nature reserve location. In contrast to earlier studies at global scale, our study did not show that nature reserves attract urbanisation. Instead, we stress that the great variety of contextual factors at the city scale makes quantitative analysis of this kind extremely difficult. However, a general neglect from planning and nature conservation agencies to recognise nature reserves’ dependence on the surrounding landscape configuration could be detrimental to sustain their values in the long-term. Hence we suggest that zones surrounding nature-protected areas should be planned and managed according to local land use history and current landscape conditions to enable and enhance necessary cross-boundary interactions.