Of niche differentiation,dispersal ability and historical legacies: what drives woody community assembly in recent Mediterranean forests?

Community assembly rules have been extensively studied, but its association with regional environmental variation and land use history remains largely unexplored. Land use history might be especially important in Mediterranean forests, considering their historical deforestation and recent afforestation. Using forest inventories and historical (1956) and recent (2000) land cover maps, we explored the following hypotheses: 1) woody species assembly is driven by environmental factors, but also by historical landscape attributes; 2) recent forests exhibit lower woody species richness than pre‐existing due to the existence of colonization credits; 3) these credits are modulated by species’ life‐forms and dispersal mechanisms. We examined the association of forest historical type (pre‐existing versus recent) with total species richness and that of diverse life‐forms and dispersal groups, also considering the effects of current environment and past landscape factors. When accounting for these effects, no significant differences in woody species richness were found between forest historical types except for vertebrate‐dispersed species. Species richness of this group was affected by the interaction of forest historical type with distance to coast and rainfall: vertebrate‐dispersed species richness increased with rainfall and distance to the coast in recent forests, while it was higher in dryer sites in pre‐existing forests. In addition, forest historical types showed differences in woody species composition associated to diverse environmental and past landscape factors. In view of these results we can conclude that: 1) community assembly in terms of species richness is fast enough to exhaust most colonization credit in recent Mediterranean forests except for vertebrate‐dispersed species; 2) for these species, colonization credit is affected by the interplay of forest history and a set of proxies of niche and landscape constraints of species dispersal and establishment; 3) woody species assemblage is mostly shaped by the species’ ecological niches in these forests.     

Soil‐based habitat partitioning in understorey palms in lower montane tropical forests

Aim Dispersal assembly and niche assembly are two competing theories proposed to explain the maintenance of species diversity in tropical forests. Dispersal theory emphasizes the role of chance colonization events and distance‐limited seed dispersal in explaining species abundance and distribution, whereas niche theory emphasizes differences among species in requirements for potentially limiting resources. Species distribution patterns in tropical forests often correlate with geology and topography, but tests of the relative importance of dispersal and niche partitioning have been hampered by an inadequate characterization of resource availability. The aim of this study was to explore how soil chemical and physical properties, climate, and geographic distance affect understorey palm communities in lower montane forests. Location Fortuna Forest Reserve, Chiriqui Province, and Palo Seco Forest Reserve, Bocas del Toro Province, in western Panama. Methods Understorey palms and soil nutrient concentrations were surveyed within 10 sites on different soil types across a 13‐km transect. Variation in palm community composition was examined in relation to spatial and environmental variables. Results The 25 understorey palm species recorded in the study were non‐randomly distributed among forests differing in soil nutrient availability. In support of dispersal theory, floristic similarity decreased predictably with increasing geographic distance. However, environmental and soil variables were also correlated with geographic distance. Floristic similarity was also highly associated with a subset of environmental variables. Variation in palm community similarity was most strongly correlated with inorganic nitrogen availability and cation concentration. A subset of soil variables had a stronger relationship with floristic similarity when geographic distance was controlled for than did geographic distance when differences in soils were controlled for. Main conclusions Both dispersal and niche processes affect palm species distribution patterns. Although spatially limited dispersal may influence species distribution patterns, soil‐based habitat associations, particularly with respect to soil nitrogen, cation availability and aluminium concentrations, remain important factors influencing palm community composition at the mesoscale level in this tropical montane forest.     

Groupings of life‐history traits are associated with distribution of forest plant species in a fragmented landscape

Questions: 1. Do relationships among forest plant traits correspond to dispersability‐persistence trade‐offs or other inter‐trait correlations found in the literature? 2. Do species groups delineated by trait similarity, differ in occurrence in ancient vs. new forests or isolated vs more continuous forest patches? 3. Are these patterns consistent for different forest types? Location: Central Belgium, near Leuven. Methods: We investigate the distributions of a large set of plant traits and combinations among all forest species occurring in patches with varying forest continuity and isolation. Through calculation of Gower's similarity index and subsequent clustering,‘emergent’ species groups are delineated. Then, the relative occurrence of these different groups in forest patches of different age and size, sustaining different forest types (alluvial vs. Quercion), and having different isolation status is compared through multivariate GLM analysis. Results: Correlations among several life history traits point towards trade‐offs of dispersability and fecundity vs. longevity. We distinguished three species groups: 1= mainly shrubs or climbers with fleshy or wind dispersed fruits and high dispersal potential; 2 = dominated by small, mainly vegetatively reproducing herbs; 3 = with spring flowering herbs with large seeds and mainly unassisted dispersal. Relative occurrence of these groups was significantly affected by forest age, area, isolation and forest type. Separate analyses for alluvial and Quercion forests indicated that the relative importance of these factors may differ, depending on forest type and species group. Both forest continuity and isolation are important in restricting the relative occurrence of forest species in alluvial forests, whatever their group membership. In Quercion forests forest patch area was the primary determinant of relative occurrence of species groups. Conclusions: It is very important to preserve the actual forest area including the spatial setting and the dispersal infrastructure within the landscape. Next, forest connectivity may be restored, but it is inherently a long process.     

Forest species in an agricultural landscape in the Netherlands: Effects of habitat fragmentation

Abstract. For 312 forest patches on sandy soils in the Netherlands, effects of fragmentation are studied of forest habitat in the past on the present occurrence of forest plant species. Using regression techniques, the numbers of forest edge, interior, zoochorous and anemochorous species, as well as occurrence of 24 individual species were related to patch area and connectivity measures. Connectivity was defined as the amount of forest habitat around patches within three zones up to 1000 m. Plant categories were distinguished by habitat type and dispersal mechanism. The results showed that number of total species and number of species of all habitat and dispersal categories increased with area. The occurrence of ten individually studied species were also positively related to area. Most of them were interior species. The number of zoochorous species increased with increasing connectivity. Also occurrence of ten individually studied species were affected by connectivity. Interior zoochorous species showed the highest percentage of affected species. The relationship of interior, animal-dispersed plants to connectivity can be explained by the limited distances covered by their dispersal agents (forest birds and ants) in a non-forest habitat. Also, some anemochorous plants appeared to be affected by connectivity, especially those with heavy seeds and potentially short distance dispersal. As not all species within a certain dispersal or habitat category react similar to area or isolation, it is suggested that differences in underlying processes of fragmentation such as local extinction and colonization need more focus.     

Rainforest expansion reduces understorey plant diversity and density in open forest of eastern Australia

The expansion of rainforest pioneer trees into long‐unburnt open forests has become increasingly widespread across high rainfall regions of Australia. Increasing tree cover can limit resource availability for understorey plant communities and reduce understorey diversity. However, it remains unclear if sclerophyll and rainforest trees differ in their competitive exclusion of understory plant communities, which contain most of the floristic diversity of open forests. Here, we examine dry open forest across contrasting fire histories (burnt and unburnt) and levels of rainforest invasion (sclerophyll or rainforest midstorey) to hindcast changes in understorey plant density, richness and composition. The influence of these treatments and other site variables (midstorey structure, midstorey composition and soil parameters) on understorey plant communities were all examined. This study is the first to demonstrate significantly greater losses of understorey species richness, particularly of dry open‐forest specialists, under an invading rainforest midstorey compared to a typical sclerophyll midstorey. Rainforest pioneers displaced over half of the understorey plant species, and reduced ground cover and density of dry forest specialists by ~90%. Significant understorey declines also occurred with increased sclerophyll midstorey cover following fire exclusion, although losses were typically less than half that of rainforest‐invaded sites over the same period. Understorey declines were closely related to leaf area index and basal area of rainforest and wattle trees, suggesting competitive exclusion through shading and potentially belowground competition for water. Around 20% of displaced species lacked any capacity for population recovery, while transient seed banks or distance‐limited dispersal may hinder recovery for a further 68%. We conclude that rainforest invasion leads to significant declines in understorey plant diversity and cover in open forests. To avoid elimination of local native plant populations in open forests, fires should occur with sufficient frequency to prevent overstorey cover from reaching a level where shade‐intolerant species fail to thrive.     

Life-history traits are correlated with geographical distribution patterns of western European forest herb species

Aim To investigate whether six plant life‐history traits that have been related to colonization ability at local scales are also related to the geographical range characteristics of 273 forest plant species. Location Continental western Europe, five countries in particular: France, Luxemburg, Belgium, the Netherlands and Germany. The region is situated between 42° and 55°N and 5°W and 15°E and has a summed total area of 971,404 km². Methods Distribution data were compiled from five national data bases and converted to a 10′ grid. Life‐history traits were taken from existing compilations of autecological information of European species. The spatial arrangement of occupied grid cells was investigated using Ripley's K. Cross‐species correlations and phylogenetically independent contrasts were used to investigate the relationships between plant life‐history traits and three range characteristics: area of occupancy, latitudinal extent and centroid latitude. Results For herbaceous species, seed dispersal mode, seed production and seed bank longevity exhibited significant associations with geographical range characteristics, including area of occupancy. Woody plant species exhibited fewer significant associations, although maximum height was positively associated with range centroid latitude within the study area. Furthermore, the ranges of species with limited dispersal ability were found to be more clustered than the ranges of species with morphological adaptations for long‐distance seed dispersal. Main conclusions For western European forest plant species, life‐history traits that are related to colonization ability at local scales are associated with variation in large‐scale geographical range characteristics. This finding implies that the distributions of some forest plant species in the study area may be limited by seed dispersal and colonization capacity rather than climate or other environmental factors.     

Colonization of recent coniferous versus deciduous forest stands by vascular plants at the local scale

Questions: 1. Are there differences among species in their preference for coniferous vs. deciduous forest? 2. Are tree and shrub species better colonizers of recent forest stands than herbaceous species? 3. Do colonization patterns of plant species groups depend on tree species composition? Location: Three deciduous and one coniferous recent forest areas in Brandenburg, NE Germany. Methods: In 34 and 21 transects in coniferous and deciduous stands, respectively, we studied the occurrence and percentage cover of vascular plants in a total of 150 plots in ancient stands, 315 in recent stands and 55 at the ecotone. Habitat preference, diaspore weight, generative dispersal potential and clonal extension were used to explain mechanisms of local migration. Regression analysis was conducted to test whether migration distance was related to species’ life‐history traits. Results: 25 species were significantly associated with ancient stands and ten species were significantly more frequent in recent stands. Tree and shrub species were good colonizers of recent coniferous and deciduous stands. In the coniferous stands, all herbaceous species showed a strong dispersal limitation during colonization, whereas in the deciduous stands generalist species may have survived in the grasslands which were present prior to afforestation. Conclusions: The fast colonization of recent stands by trees and shrubs can be explained by their effective dispersal via wind and animals. This, and the comparably efficient migration of herbaceous forest specialists into recent coniferous stands, implies that the conversion of coniferous into deciduous stands adjacent to ancient deciduous forests is promising even without planting of trees.     

Colonization front of the understorey palm Astrocaryum sciophilum in a pristine rain forest of French Guiana

Aims Astrocaryum sciophilum (Miq.) Pulle (Arecaceae) is an understorey palm, endemic to north‐eastern South America with a patchy distribution. We tested the hypothesis that the spatial distribution of this palm species is not in equilibrium but is slowly colonizing the forest understorey. Location Inventories and seed dispersal studies were conducted in the undisturbed tropical forest close to the Nouragues research station, French Guiana. Additional data were collected in the entire territory of French Guiana. Methods We studied the demography of A. sciophilum on a 20‐ha plot located at the edge of its distribution. The age of the palms was estimated by postulating an exponentially decreasing abundance by age class. Direct seed dispersal experiments were also conducted, to estimate dispersal parameters. The seeds of A. sciophilum were dispersed only by rodents. This information was used to parameterize a forest growth simulator, to study the spatial spread of this species. Results Within the sampling plot, the density of A. sciophilum dropped sharply from about 500 individuals per hectare to zero. The maturation age was estimated to be 170 ± 70 years, and over 55 years with 95% confidence. Seed‐dispersal experiments yielded an average seed dispersal distance of 11 m and a maximum estimated dispersal distance of 125 m across a generational span of 55 years to maturity. Therefore, the maximal estimated colonization speed is 2.3 m/y. Conclusions Empirical results and numerical simulations suggest that the boundary of the A. sciophilum population is a colonization front, and that the range of this species is slowly expanding. The implications of this result in respect of palaeoenvironmental changes in this region are discussed.     

Seed dispersal mechanisms and the vegetation of forest islands in a West African forest-savanna mosaic (Comoé National Park,Ivory Coast)

The transition zone between forest and savanna is typically characterized by a dynamic patchwork of forest and savanna. We studied the woody plant species composition of 49 forest islands, 18 savanna, and 3 gallery forest plots in the Comoé National Park (Ivory Coast), West Africa's largest savanna reserve. TWINSPAN makes a clear distinction in vegetational composition between these three major habitat types but, nevertheless, more than 50% of the 292 species occur in at least 2 of them. The gallery forest is dominated by Cynometra megalophylla (Caesalpiniaceae), a genus known to dominate humid forests in other afrotropical regions. Ordination reveals four distinct categories of forest islands, (1) humid forests comparable to the gallery forest in their species composition, (2) dry disturbed and (3) dry undisturbed forests and (4) forests formerly inhabited by humans. Disturbed forests harbor more savanna species but also a distinct group of disturbance-tolerant forest species. Compared to other forest species, in this latter group we found an exceptionally high fraction of animal dispersed species (80% vs. 58%), while wind dispersed species or species lacking long distance seed dispersal mechanisms were correspondingly rare. This pattern occurs in spite of the fact that the frequency of wind dispersed species in general increases from dense humid forests to open dry forests and savanna. Species lacking long distance dispersal mechanisms are most abundant among those specialized on humid forests. These observations suggest that the species composition of forest islands is to some extent determined by the seed dispersal abilities of the different species.     

Diversity–invasibility relationships across multiple scales in disturbed forest understoreys

Non-native plant species richness may be either negatively or positively correlated with native species due to differences in resource availability, propagule pressure or the scale of vegetation sampling. We investigated the relationships between these factors and both native and non-native plant species at 12 mainland and island forested sites in southeastern Ontario, Canada. In general, the presence of non-native species was limited: <20% of all species at a site were non-native and non-native species cover was <4% m⁻² at 11 of the 12 sites. Non-native species were always positively correlated with native species, regardless of spatial scale and whether islands were sampled. Additionally, islands had a greater abundance of non-native species. Non-native species richness across mainland sites was significantly negatively correlated with mean shape index, a measure of the ratio of forest edge to area, and positively correlated with the mean distance to the nearest forest patch. Other factors associated with disturbance and propagule pressure in northeastern North America forests, including human land use, white-tailed deer populations, understorey light, and soil nitrogen, did not explain non-native richness nor cover better than the null models. Our results suggest that management strategies for controlling non-native plant invasions should aim to reduce the propagule pressure associated with human activities, and maximize the connectivity of forest habitats to benefit more poorly dispersed native species.     

The comparative importance of species traits and introduction characteristics in tropical plant invasions

Aim We used alien plant species introduced to a botanic garden to investigate the relative importance of species traits (leaf traits, dispersal syndrome) and introduction characteristics (propagule pressure, residence time and distance to forest) in explaining establishment success in surrounding tropical forest. We also used invasion scores from a weed risk assessment protocol as an independent measure of invasion risk and assessed differences in variables between high‐ and low‐risk species. Location East Usambara mountains, Tanzania. Methods Forest transect surveys identified species establishing in disturbed and intact forest. Leaf traits (specific leaf area and foliar nutrient concentrations) were measured from leaves sampled in high‐light environments. Results A leaf traits spectrum was apparent, but species succeeding or failing to establish in either disturbed or intact forest were not located in different parts of the spectrum. Species with high invasion risk did not differ in their location on the leaf trait spectrum compared with low‐risk species but were more likely to be bird/primate‐dispersed. For 15 species establishing in forest quadrats, median canopy cover of quadrats where seedlings were present was correlated with a species value along the leaf trait spectrum. Species establishing in disturbed forest were planted in twice as many plantations and were marginally more likely to be bird‐ or primate‐dispersed than species failing to become established in disturbed forest. Establishment in intact forest was more likely for species planted closer to forest edges. Main conclusions Leaf and dispersal traits appear less important in the colonization of tropical forest than introduction characteristics. It appears, given sufficient propagule pressure or proximity to forest, alien species are much more likely to establish independently of leaf traits or dispersal syndrome in continental tropical forests.     

Legacies of the past in the present-day forest biodiversity: a review of past land-use effects on forest plant species composition and diversity

Particularly in the temperate climate zone many forests have, at some moment in their history, been used as agriculture land. Forest cover is therefore often not as stable as it might look. How forest plant communities recovered after agriculture was abandoned allows us to explore some universal questions on how dispersal and environment limit plant species abundance and distribution. All studies looking at the effects of historical land use rely on adequate land use reconstruction. A variety of tools from maps, archival studies, and interviews to field evidence and soil analyses contribute to that. They allow us to distinguish ancient from recent forests and many studies found pronounced differences in forest plant species composition between them. A considerable percentage of our forest flora is associated with ancient forests. These ancient forest plant species (AFS) all have a low colonization capacity, suggesting that dispersal in space (distance related) and time (seed bank related) limit their distribution and abundance. However recent forests generally are suitable for the recruitment of AFS. There is clear evidence that dispersal limitation is more important than recruitment limitation in the distribution of AFS. Dispersal in time, through persistent seed banks, does not play a significant role. Ancient forests are not necessary more species-rich than recent forest, but if diversity is limited to typical forest plant species then ancient forests do have the highest number of plant species, making them highly important for nature conservation. The use of molecular markers, integrated approaches and modelling are all part of the way forward in this field of historical ecology.     

Species and structural diversity of church forests in a fragmented Ethiopian Highland landscape

Question: Thousands of small isolated forest fragments remain around churches (“church forests”) in the almost completely deforested Ethiopian Highlands. We questioned how the forest structure and composition varied with altitude, forest area and human influence. Location: South Gondar, Amhara National Regional State, Northern Ethiopia. Methods: The structure and species composition was assessed for 810 plots in 28 church forests. All woody plants were inventoried, identified and measured (stem diameter) in seven to 56 10 m x 10‐m plots per forest. Results: In total, 168 woody species were recorded, of which 160 were indigeneous. The basal area decreased with tree harvest intensity; understorey and middle‐storey density (<5 cm DBH trees) decreased with grazing; overstorey density (>5 cm DBH trees) increased with altitude. The dominance of a small set of species increased with altitude and grazing intensity. Species richness decreased with altitude, mainly due to variation in the richness of the overstorey community. Moreover, species richness in the understorey decreased with grazing intensity. Conclusions: We show how tree harvesting intensity, grazing intensity and altitude contribute to observed variations in forest structure, composition and species richness. Species richness was, however, not related to forest area. Our study emphasizes the significant role played by the remaining church forests for conservation of woody plant species in North Ethiopian Highlands, and the need to protect these forests for plant species conservation purposes.     

Restoration of the understorey layer of recent forest bordering ancient forest

Abstract. Recently established forests are commonly characterized by an impoverished understorey. Restoration is mostly based on spontaneous secondary succession, but little is known about the time period needed to achieve a community species pool with species composition equal to that of ancient forests. Vegetation in transects of 197 plots in 13 recent forest stands contiguous to the Meerdaalwoud ancient forest complex was surveyed. The recent forest stands ranged in age from 36 to 132 yr. The community species pool was described with an ecological, functional and phytosociological approach and based on groups derived from a CCA. Differences in community species pool between age classes of recent forest stands were analysed. During establishment of a new forest competitive species, forest edge species and species with high Ellenberg values for light and nitrogen and a more persistent seed bank will dominate the understorey. After 90 yr of succession the cover by these species decreases and reaches equal values to ancient forest after ca. 105 yr. A large number of forest species will be able to colonize the forest in less than 90 yr. Some typical forest species, however, have very low colonization rates and still have low cover in recent forest more than 105 yr old, so that complete restoration of the understorey requires a time period of over a century. Anthropogenic introduction of forest plant species may reduce the time required for ancient forest vegetation equality.     

Do forest plants conform to the theory of island biogeography: the case study of bog islands

Biodiversity conservation is confronted with increasing risk of extinction in isolated small-area remnants and the limitation of species to colonize recently formed habitats. We hypothesized that the equilibrium pattern of forest herb layer in long-term fragmented landscape should comply with the theory of island biogeography. Forests on mineral soil islands located in large mires of western Estonia were considered as dispersal target habitats, and forests on mainland and peninsulas in mires as sources. Species richness was the lowest in mainland forests and the effect was confounded by habitat structure, suggesting a negative effect of silvicultural management in easily accessible forests. We observed the ‘small island effect’, i.e. greater overall species richness in small-area habitats, which was determined by the habitat preference of shade tolerant generalists. The average species richness of common mainland forest specialists varied little, but capitalizing on the traditional approach and analyzing only island data, weak effects of distance and habitat quality were detected. At single species level, unexpectedly, many habitat specialists were observed to have successfully dispersed to islands, indicating insufficient knowledge of the long-distance dispersal mechanisms of forest-dwelling plants. In fragmented forest landscapes the theory of island biogeography can be applied to habitat specialist plant species, but only regarding the effect of isolation and in conditions of persistent forest structural quality. In the light of global changes, optimized conservation planning should primarily target on (i) the conservation of ancient habitat fragments independent of their current area, and (ii) the promotion of diversity of potential dispersal vectors in the landscape.     

Lichen colonization patterns show minor effects of dispersal distance at landscape scale

The role of dispersal in controlling the distribution of species at landscape scale (10²–10⁴ m) is still a matter of dispute. Here, we use the early colonization pattern of 23 epiphytic lichen species in a former tree‐less heathland landscape (170 km²) to test three hypotheses on how a landscape is colonized: A) mainly by long‐distance dispersal (LDD), B) by rare LDD events followed by limited local dispersal, and C) mainly by limited dispersal, resulting in a colonization front. The study system consisted of a chronosequence of 94 habitat patches constituting 0.4% of the landscape area, with a minimum inter‐site distance of 0.2 km. We used generalized linear mixed models with Bayesian inference to test predictions from the hypotheses. When age of sites and habitat area were accounted for, additional effects of geographical position of sites (distance from old sites, distance‐dependent relative propagule pressure, and distance from border of study area) on the probability of colonization by lichen species were small. Furthermore, species richness of sites did not depend on geographical position, either. Our results support a colonization process mainly governed by LDD at landscape scale, and that local stepwise colonization was not important. We argue that passively dispersed species with numerous small propagules tend to exhibit patchy populations with extensive dispersal at the landscape scale, rather than behaving like classical metapopulations.     

Effects of tree species composition on within‐forest distribution of understorey species

Question: Do tree species, with different litter qualities, affect the within‐forest distribution of forest understorey species on intermediate to base‐rich soils? Since habitat loss and fragmentation have caused ancient forest species to decline, those species are the main focus of this study. Location: Three ancient forests, along a soil gradient from acidification‐sensitive to base‐rich, were studied: Limbrichterbosch and Savelsbos in The Netherlands and Holtkrat in Denmark. Methods: Canopy and soil surveys along transects generated data for Redundancy Analysis on tree – humus relationships. We analysed the distribution of forest plant species with Canonical Correspondence Analysis. The explanatory factors were soil characteristics (pH, organic matter, loam content and thickness of the humus layers), external crown projection, ground water and canopy data. We further analysed the relationship between forest species and humus characteristics with Spearman correlations. Results: Tree species have a significant impact on humus characteristics through the nature of their litter. Humus characteristics significantly explain the distribution of forest understorey species. The pH of the first 25 cm mineral soil and the thickness of the F‐ (fermentation) layer are the primary factors affecting the distribution of ancient forest species. Conclusion: This study indicates that the species composition of the forest canopy affects the distribution of forest understorey species. Ancient forest species are more abundant and frequent underneath trees with base‐rich litter. On acidification‐sensitive soils these relationships were stronger than on more base‐rich, loamy soils.     

Impact of the spatial uncertainty of seed dispersal on tree colonization dynamics in a temperate forest

An aggregated distribution of dispersed seeds may influence the colonization process in tree communities via inflated spatial uncertainty. To evaluate this possibility, we studied 10 tree species in a temperate forest: one primarily barochorous, six anemochorous and two endozoochorous species. A statistical model was developed by combining an empirical seed dispersal kernel with a gamma distribution of seedfall density, with parameters that vary with distance. In the probability density, the fitted models showed that seeds of Fagaceae (primarily barochorous) and Betulaceae (anemochorous) were disseminated locally (i.e. within 60 m of a mother tree), whereas seeds of Acer (anemochorous) and endozoochorous species were transported farther. Greater fecundity compensated for the lower probability of seed dispersal over long distances for some species. Spatial uncertainty in seedfall density was much greater within 60 m of a mother tree than farther away, irrespective of dispersal mode, suggesting that seed dispersal is particularly aggregated in the vicinity of mother trees. Simulation results suggested that such seed dispersal patterns could lead to sites in the vicinity of a tree being occupied by other species that disperse seeds from far away. We speculate that this process could promote coexistence by making the colonization rates of the species more similar on average and equalizing species fitness in this temperate forest community.     

The role of landscape configuration in plant composition of floodplain forests across different physiographic areas

Questions: What is the relative importance of landscape variables compared to habitat quality variables in determining species composition in floodplain forests across different physiographic areas? How do species composition and species traits relate to effects of particular landscape variables? Do lowland and mountain areas differ in effects of landscape variables on species composition? Location: Southern Czech Republic. Methods: A total of 240 vegetation relevés of floodplain forests with measured site conditions were recorded across six physiographic areas. I tested how physiographic area, habitat quality variables and landscape variables such as current land‐cover categories, forest continuity, forest size and urbanization influenced plant species composition. I also compared how mountain and lowland areas differ in terms of the relative importance of these variables. To determine how landscape configuration affects the distribution of species traits, relationships of traits and species affinity with landscape variables were tested. Results: Among landscape variables, forest continuity, landscape forest cover and distance to nearest settlement altered the vegetation. These variables also influenced the distributions of species traits, i.e. life forms, life strategies, affinity to forest, dispersal modes, seed characteristics, flooding tolerance and Ellenberg indicator values for nitrogen, light, moisture and soil reaction. Nevertheless, physiographic area and habitat quality variables explained more variation in species composition. Landscape variables were more important in lowland areas. Forest continuity affected species composition only in lowlands. Conclusions: Although habitat quality and physiographic area explained more vegetation variability, landscape configuration was also a key factor influencing species composition and distribution of species traits. However, the results are dependent on forest geographical location, with lowland forests being more influenced by landscape variables compared to mountain forests.