The effect of landscape fragmentation and forest continuity on forest floor species in two regions of Denmark

Abstract. Forest species composition was recorded in 82 forests in the Himmerland and Hornsherred regions in Denmark and analysed with respect to isolation (distance to other forests and areas of forest), forest continuity (older or younger than 200 yr), soil pH, tree species composition and seed dispersal groups. Continuity and isolation measures were correlated with forest species richness in Hornsherred. Myrmecochorous, autochorous, anemoballistic and endozoochorous species were markedly fewer in recent than in ancient forests. In Himmerland, patterns were much weaker and few significant correlations were found between forest species richness or different seed dispersal groups and continuity or isolation of the forests. Differences between the two regions may result from less intensive land use, a more humid climate and a smaller species pool with less species with short distance dispersal in Himmerland. Landscape fragmentation therefore appears to limit forest species’recolonization more in Hornsherred than in Himmerland.     

Dispersal and seed limitation affect diversity and productivity of montane grasslands

So far, seed limitation as a local process, and dispersal limitation as a regional process have been largely neglected in biodiversity–ecosystem functioning research. However, these processes can influence both local plant species diversity and ecosystem processes, such as biomass production. We added seeds of 60 species from the regional species pool to grassland communities at 20 montane grassland sites in Germany. In these sites, plant species diversity ranged from 10 to 34 species m⁻² and, before manipulation, diversity was not related to aboveground biomass, which ranged from 108 to 687 g m⁻². One year after seed addition, local plant species richness had increased on average by six species m⁻² (29%) compared with control plots, and this increase was highest in grasslands with intermediate productivity. The increased diversity after adding seeds was associated with an average increase of aboveground biomass of 36 g m⁻² (14.8%) compared with control plots. Thus, our results demonstrate that a positive relationship between changes in species richness and productivity, as previously reported from experimental plant communities, also holds for natural grassland ecosystems. Our results show that local plant communities are dispersal limited and a hump‐shaped model appears to be the limiting outline of the natural diversity–productivity relationship. Hence, the effects of dispersal on local diversity can substantially affect the functioning of natural ecosystems.     

Predictability of plant species composition from environmental conditions is constrained by dispersal limitation

Despite recent modelling approaches integrating the effects of niche-based processes and dispersal-based processes on local plant species composition, their relative importance is still not clear. We test whether the predictability of local species composition from environmental conditions is influenced by dispersal traits. We analyzed a large database with co-occurrence data, using ordination techniques (DCA and CCA) to identify the major environmental determinants of species composition. The percentage of explained variance in occurrence was quantified for individual species with CCA. Effects of life-history traits on the predictability of occurrence patterns were tested by means of regression analysis, using a generalized linear models approach. The results reveal close correlations between species composition and environmental conditions, implying that the predictability of the set of species that might occur in a given environmental setting ("habitat species pool") is high. The habitat species pool, however, reflects the potential species composition, and not the actual local situation. At the level of individual species, a large proportion (>90%) of the variation in occurrence remained unexplained. Predictability of species occurrence patterns was increased by a greater capacity for long-distance dispersal, greater adult longevity and the capacity to build a persistent seed bank. The results indicate that the predictability of species composition from environmental conditions is reduced by a few orders of magnitude by dispersal limitation and that poor dispersers are underrepresented.     

Evidence for community assembly constraints during succession in dune slack plant communities

Community assembly during succession can be constrained by both local and regional factors. Despite an increasing regional species pool size during succession, we found a limit on the number of species in 1 × 1 m plots in dune slacks. Three alternative hypotheses (habitat heterogeneity, dispersal limitation and niche limitation) explaining this community saturation were tested. A null model analysis showed that species richness in the plots had an unusually low variance suggesting that beta habitat diversity was not likely to explain the limitation on species richness. Because we did not find a correlation between the distribution of species over the slack and their dispersal capacity, we also excluded the dispersal limitation hypothesis. Finally, a guild proportionality analysis revealed that the abundances of forb, graminoid and ruderal species showed low an unusually low variance over all age classes involved. This provides evidence for nonrandom community assembly during succession, likely to be determined by competitive exclusion among species of the same guild.     

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.     

Pervasive effects of dispersal limitation on within- and among-community species richness in agricultural landscapes

Aim To determine whether the effect of habitat fragmentation and habitat heterogeneity on species richness at different spatial scales depends on the dispersal ability of the species assemblages and if this results in nested species assemblages. Location Agricultural landscapes distributed over seven temperate Europe countries covering a range from France to Estonia. Methods We sampled 16 local communities in each of 24 agricultural landscapes (16 km²) that differ in the amount and heterogeneity of semi‐natural habitat patches. Carabid beetles were used as model organisms as dispersal ability can easily be assessed on morphological traits. The proximity and heterogeneity of semi‐natural patches within the landscape were related to average local (alpha), between local (beta) and landscape (gamma) species richness and compared among four guilds that differ in dispersal ability. Results For species assemblages with low dispersal ability, local diversity increased as the proximity of semi‐natural habitat increased, while mobile species showed an opposite trend. Beta diversity decreased equally for all dispersal classes in relation to proximity, suggesting a homogenizing effect of increased patch isolation. In contrast, habitat diversity of the semi‐natural patches affected beta diversity positively only for less mobile species, probably due to the low dispersal ability of specialist species. Species with low mobility that persisted in highly fragmented landscapes were consistently present in less fragmented ones, resulting in nested assemblages for this mobility class only. Main conclusions The incorporation of dispersal ability reveals that only local species assemblages with low dispersal ability show a decrease of richness as a result of fragmentation. This local species loss is compensated at least in part by an increase in species with high dispersal ability, which obscures the effect of fragmentation when investigated across dispersal groups. Conversely, fragmentation homogenizes the landscape fauna for all dispersal groups, which indicates the invasion of non‐crop habitats by similar good dispersers across the whole landscape. Given that recolonization of low dispersers is unlikely, depletion of these species in modern agricultural landscapes appears temporally pervasive.     

Seed arrival, ecological filters, and plant species richness: a meta-analysis

Theoretical models predict that effects of dispersal on local biodiversity are influenced by the size and composition of the species pool, as well as ecological filters that limit local species membership. We tested these predictions by conducting a meta-analysis of 28 studies encompassing 62 experiments examining effects of propagule supply (seed arrival) on plant species richness under contrasting intensities of ecological filters (owing to disturbance and resource availability). Seed arrival increased local species richness in a wide range of communities (forest, grassland, montane, savanna, wetland), resulting in a positive mean effect size across experiments. Mean effect size was 70% higher in disturbed relative to undisturbed communities, suggesting that disturbance increases recruitment opportunities for immigrating species. In contrast, effect size was not significantly influenced by nutrient or water availability. Among seed-addition experiments, effect size was positively correlated with species and functional diversity within the pool of added seeds (species evenness and seed-size diversity), primarily in disturbed communities. Our analysis provides experimental support for the general hypothesis that species pools and local environmental heterogeneity interactively structure plant communities. We highlight empirical gaps that can be addressed by future experiments and discuss implications for community assembly, species coexistence, and the maintenance of biodiversity.     

Spatial context strongly affects community composition of both passively and actively dispersing pool invertebrates in a highly heterogeneous landscape

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Factors influencing species richness in lacustrine zooplankton

Frequent dispersal events are expected to elevate local species richness in island-like habitats such as lakes. However, the importance of dispersal can be hard to evaluate if other factors cause large background variability in species composition and richness. In this paper, we review empirical studies on ecological factors known or expected to influence species richness in zooplankton communities of inland lakes. We then present summaries of two recent case studies. Our objectives are twofold: we first look for effects of biotic interactions on species richness and species composition, and then evaluate whether the expected effects of dispersal are likely to be detected on a background of large variability caused by other ecological factors and interactions. Species richness within lakes appears to be primarily controlled by factors related to lake size, lake productivity, water quality, and fish predation levels. One case study indicated a slight, but significant, positive effect of lake density and lake area in the surrounding landscape on species richness, suggesting that frequent dispersal events may enhance species richness. This local variation in species richness is superimposed on regional variation in species pools.     

Large-scale spatial ecology of dung beetles

Recent modelling work shows that the composition of local communities can be influenced by the configuration of the surrounding landscape, but many of these models assume that all community members display the same type of extinction‐colonization dynamics. I use Aphodius dung beetles to test the hypothesis that interspecific differences in habitat selection and dispersal capacity may translate into differences in spatial population dynamics, even among closely related species coexisting on the same resource. If this is true, then groups of species with different characteristics would show different responses to landscape configuration. I first divided the area of Finland into a grid, and used collection records to describe regional variation in the Aphodius fauna of open cattle pastures. I then sampled dung beetles on 131 cattle farms, to examine whether the subset of species found on each farm was related to the density of pastures in the surrounding grid square. Finally, I used historical records to analyze changes in dung beetle communities during the last century, when there was great loss of pasture. Overall, I found no relationship between landscape characteristics and the total proportion of the regional species pool that was found on each farm. However, the distribution of species among guilds with different habitat specificity did relate to the configuration of the landscape, and the pattern was most pronounced in a specialists species with limited dispersal. Associations between community structure and landscape configuration were superimposed on two much larger and stronger patterns: a large‐scale latitudinal gradient in regional species richness, and a decelerating gain of species to local communities with an increasing regional species pool. I conclude that ecological variation among community members is a crucial factor in the analysis of local community composition, and that local species richness should always be conditioned on regional richness.     

The role of historical processes in determining tree species richness in the forests of Western Caucasus

To estimate the role of history in variation of tree species richness in the forests of the Western Caucasus we analyzed correlation between their local richness (S--the mean number of species per 300 m2) and size of actual species pool (N--the number of species per 1 ha). If compared communities are differently distant from the point of evolutionary equilibrium one should expect a significant variation in correlation between S and N (determined with the greater sensitivity of N than S in respect of historical factors). The lower value of N/S corresponds to less saturated level of historically determined species richness. A mean N/S ratio in Japana temperate broadleaved forests (Masaki et al., 1999) provided the basis for analysis. The present tree species richness of the forest communities in the 1 ha plots seem essentially determined by the historical processes. The mountain forest communities of Western Caucasus are characterized on the average with lower saturation level of the actual tree species pool in comparison with the Japan temperate broad-leaved forest communities. On the Western Caucasus the middle mountain beech and coniferous-broadleaved communities (400-1600 m a.s.l.) are characterized with the higher saturation level of the actual species pool in comparison with communities located lower and higher. These results confirm published historical reconstructions, according to which the middle mountain forest communities in the Western Caucasus are older than forests located higher or lower. Present low mountain forests of the southern (to Black Sea) and the northern macroslopes of the Western Caucasus are characterized with similar saturation level of the actual species pool. These data agree with the assumption of Dolukhanov (1980) that low mountain zone of the southern macroslope was not a refuge for tree species in Pleistocene.     

Which functional processes control the short-term effect of grazing on net primary production in grasslands?

For unicellular organisms, a lack of effects of local species richness on ecosystem function has been proposed due to their locally high species richness and their ubiquitous distribution. High dispersal ability and high individual numbers may enable unicellular taxa to occur everywhere. Using our own and published data sets on uni- and multicellular organisms, we conducted thorough statistical analyses to test whether (1) unicellular taxa show higher relative local species richness compared to multicellular taxa, (2) unicellular taxa show lower slopes of the species:area relationships and species:individuals relationships, and (3) the species composition of unicellular taxa is less influenced by geographic distance compared to multicellular taxa. We found higher local species richness compared to the global species pool for unicellular organisms than for metazoan taxa. The difference was significant if global species richness was conservatively estimated but not if extrapolated, and therefore higher richness estimates were used. Both microalgae and protozoans showed lower slopes between species richness and sample size (area or individuals) compared to macrozoobenthos, also indicating higher local species richness for unicellular taxa. The similarity of species composition of both benthic diatoms and ciliates decreased with increasing geographic distance. This indicated restricted dispersal ability of protists and the absence of ubiquity. However, a steeper slope between similarity and distance was found for polychaetes and corals, suggesting a stronger effect of distance on the dispersal of metazoans compared to unicellular taxa. In conclusion, we found partly different species richness patterns among uni- and multicellular eukaryotes, but no strict ubiquity of unicellular taxa. Therefore, the effect of local unicellular species richness on ecosystem function has to be reanalyzed. Macroecological patterns suggested for multicellular organisms may differ in unicellular communities.     

Succession of Ephemeral Secondary Forests and Their Limited Role for the Conservation of Floristic Diversity in a Human-Modified Tropical Landscape

Both local- and landscape-scale processes drive succession of secondary forests in human-modified tropical landscapes. Nonetheless, until recently successional changes in composition and diversity have been predominantly studied at the patch level. Here, we used a unique dataset with 45 randomly selected sites across a mixed-use tropical landscape in central Panama to study forest succession simultaneously on local and landscape scales and across both life stages (seedling, sapling, juvenile and adult trees) and life forms (shrubs, trees, lianas, and palms). To understand the potential of these secondary forests to conserve tree species diversity, we also evaluated the diversity of species that can persist as viable metapopulations in a dynamic patchwork of short-lived successional forests, using different assumptions about the average relative size at reproductive maturity. We found a deterministic shift in the diversity and composition of the local plant communities as well as the metacommunity, driven by variation in the rate at which species recruited into and disappeared from the secondary forests across the landscape. Our results indicate that dispersal limitation and the successional niche operate simultaneously and shape successional dynamics of the metacommunity of these early secondary forests. A high diversity of plant species across the metacommunity of early secondary forests shows a potential for restoration of diverse forests through natural succession, when trees and fragments of older forests are maintained in the agricultural matrix and land is abandoned or set aside for a long period of time. On the other hand, during the first 32 years the number of species with mature-sized individuals was a relatively small and strongly biased sub-sample of the total species pool. This implies that ephemeral secondary forests have a limited role in the long-term conservation of tree species diversity in human-modified tropical landscapes.     

Size of the local species pool determines invasibility of a C4-dominated grassland

The size of the local species pool (i.e., species surrounding a community capable of dispersal into that community) and other dispersal limitations strongly influence native plant community composition. However, the role that the local species pool plays in determining the invasibility of communities by exotic plants remains to be evaluated. We hypothesized that the richness and abundance of exotic species would be greater in C4‐dominated grassland communities if the local species pool included a larger proportion of exotic species. We also predicted that an increase in the exotic species pool would increase the invasibility of sites thought to be resistant to invasion (annually burned grassland). To test these hypotheses, study plots were established within two long‐term (>20 yr) fire experiments at a tallgrass prairie preserve in NE Kansas (USA). Study plots were surrounded by either a small pool of exotic species (small species pool (SSP) plots; six species) or a larger exotic species pool (large species pool (LSP) plots; 18 species). We found that richness and absolute cover of exotic species was significantly (P<0.001) lower (～70 and 90%, respectively) in annually burned compared to unburned plots, regardless of the size of the exotic species pool. As predicted, exotic species richness was higher (P<0.001) for LSP plots (3.9 per 250 m2) than for SSP plots (0.7 per 250 m2); however, absolute cover was unaffected by the size of the exotic species pool. In the absence of fire, plots with a LSP had four times as many exotic species than SSP plots. An increase in the local exotic species pool also increased the invasibility of annually burned grassland. Indeed, richness of exotic plant species in annually burned LSP plots did not differ from unburned plots with a SSP, indicating that a larger pool of exotic species countered the negative effects of fire. These findings have important implications for predicting how the invasion of plant communities may respond to human‐induced global changes, such as habitat fragmentation. Community characteristics or factors such as frequent fires in grasslands may impart resistance to invasions by exotic species in large, intact ecosystems. However, when a large pool of exotic species is present, frequent fire may not be sufficient to limit the invasions of exotic plants in fragmented landscapes.     

The species pool concept as a framework for studying patterns of plant diversity

Co‐existence theories fail to adequately explain observed community patterns (diversity and composition) because they mainly address local extinctions. For a more complete understanding, the regional processes responsible for species formation and geographic dispersal should also be considered. The species pool concept holds that local variation in community patterns is dependent primarily on the availability of species, which is driven by historical diversification and dispersal at continental and landscape scales. However, empirical evidence of historical effects is limited. This slow progress can be attributed to methodological difficulties in determining the characteristics of historical species pools and how they contributed to diversity patterns in contemporary landscapes. A role of landscape‐scale dispersal limitation in determining local community patterns has been demonstrated by numerous seed addition experiments. However, disentangling general patterns of dispersal limitation in communities still requires attention. Distinguishing habitat‐specific species pools can help to meet both applied and theoretical challenges. In conservation biology, the use of absolute richness may be uninformative because the size of species pools varies between habitats. For characterizing the dynamic state of individual communities, biodiversity relative to species pools provides a balanced way of assessing communities in different habitats. Information about species pools may also be useful when studying community assembly rules, because it enables a possible mechanism of trait convergence (habitat filtering) to be explicitly assessed. Empirical study of the role of historic effects and dispersal on local community patterns has often been restricted due to methodological difficulties in determining habitat‐specific species pools. However, accumulating distributional, ecological and phylogenetic information, as well as use of appropriate model systems (e.g. archipelagos with known biogeographic histories) will allow the species pool concept to be applied effectively in future research.     

When do localized natural enemies increase species richness?

The Janzen–Connell hypothesis states that local species‐specific density dependence, mediated through specialist enemies of offspring such as fungal pathogens and insect seed predators, can facilitate coexistence of species by preventing recruitment near conspecific adults. We use spatially explicit simulation models and analytical approximations to evaluate how spatial scales of offspring and enemy dispersal affect species richness. In comparison with model communities in which both offspring and enemies disperse long distances, species richness is substantially decreased when offspring disperse long distances and enemies disperse short distances. In contrast, when both offspring and enemies disperse short distances species richness more than doubles and adults of each species are highly spatially clumped. For the range of conditions typical of tropical forests, locally dispersing specialist enemies may decrease species richness relative to enemies that disperse long distances. In communities where dispersal distances of both offspring and enemies are short, local effects may enhance species richness.     

The species pool hypothesis: the necessity to shift emphasis

The species pool of a biological community is determined as a group of species that inhabit some area and potentially can be included in a given community. The species pool hypothesis, i.e. the assumption that the size of species pool strongly influences species richness of local community can be confirmed if there is positive linear relationship between these two variables. The results of hypothesis testing however are not obvious. For example, correlation between local richness and species pool size can be caused by their dependence on the third variable--capacity of environment. It seems that in case of decreasing area occupied by local community the environmental conditions become more important than species pool size. If that is true, the influence of species pool on local species richness is not significant. However one can estimate the degree of unsaturation of species pool on the basis of relationships between the number of species in small locations occupied by similar local communities and their species pool. We think, that study of local and regional species richness should shift the emphasis--from the analysis of species pool influence on local community richness to the estimation of historical, ecological and anthropogenic factors in variation of species pool size. The local species richness should be considered rather as a tool (allowing to compare the species capacity of biological communities), than as an object of such study.     

Seed dispersal modes of tree species and their relation with altitudinal gradient from tropical forests of Eastern Ghats,India

The present study reports the seed dispersal modes of tree species and their relationship with altitudinal gradient from six tropical forests of Eastern Ghats located in India. Anemochory, autochory and zoochory are the three major types of seed dispersal modes recognized for the 272 tree species recorded from a total of 120 0.5 ha area. The predominant dispersal mode is zoochory with 58% of the total tree abundance and 69% of total species richness, followed by autochory with 30% and 19%, and anemochory 12% and 12%, respectively. One way ANOVA revealed that the species richness as well as tree abundance of the three dispersal modes varied significantly across the six tropical forest sites. Regression analysis revealed that both the tree species richness and abundance of zoochory dispersal mode had positive relation with altitude of forest location, while, no such relation was observed for the other two dispersal modes. This study shows that the zoochory increases in species richness and tree abundance with increasing altitude. This may be attributed to high rainfall supporting evergreen forest types with high ratio of animal dispersed tree species in forests at high altitude. However, further studies on these aspects are warranted to draw a significant statement. The predomination of zoochory dispersal mode in this study is comparable with earlier studies in tropical forests elsewhere.     

Niche assembly of epiphytic bryophyte communities in the Guianas: a regional approach

Aim Epiphytic bryophyte communities of tropical forests show a gradient in species composition from the base to the top of the host trees, indicating a strong role of niche assembly. This pattern, however, has never been tested at a regional scale. The aim of this study was to test whether niche assembly, rather than dispersal limitation, predominantly drives species composition of bryophyte communities across large spatial scales. Location Three lowland forests in the Guianas: one near Saul, French Guiana; and two near Mabura Hill, Guyana. Methods Communities of epiphytic bryophytes were sampled from six different height zones of several trees in three lowland forests. We analysed the composition of these communities using detrended correspondence analysis in order to find the best explanatory variable for the variation in community composition. A multi‐response permutation procedure was used to test the significance of grouping communities by height zone. We conducted an indicator species analysis to classify species as specialists or generalists and then tested, through weighted averaging, if specialists would indeed maintain their preferred height zone across the Guianas. Results Community composition was explained mainly by height zone. The similarity among communities inhabiting the same height zone of trees, across a distance of up to 640 km, was higher than the similarity among communities established along the vertical gradient of a single standing tree (30–50 m). More than half (57%) of the species had a preferred height zone, and the preference was consistent: species occupied roughly the same height zone on host trees in the different localities. The three local communities investigated were found to belong to the same regional species pool. Main conclusions Throughout the Guianas, epiphytic bryophyte communities are drawn from the same regional species pool, and their composition is shaped by micro‐environmental conditions. The predominance of niche assembly over dispersal assembly rules is consistently found at both local and regional scales.     

Travelling to a former sea floor: colonization of forests by understorey plant species on land recently reclaimed from the sea

Questions: What are important forest characteristics determining colonization of forest patches by forest understorey species? Location: Planted forests on land recently reclaimed from the sea, the Netherlands. Methods: We related the distribution of forest specialist species in the understorey of 55 forests in Dutch IJsselmeer polders to the following forest characteristics: age, area, connectivity, distance to mainland (as a proxy for distance to seed source) and path density. We used species of the Fraxino‐Ulmetum association for the Netherlands as reference for species that could potentially occur in the study area. Results: Area and age of the surveyed forests explained a large part of the variation in overall species composition and species number of forest plant species. The importance of connectivity and distance to the mainland of forest habitats became apparent only at a more detailed level of dispersal groups and individual species. The importance of forest parameters differed between dispersal groups and also between individual species. After 60 years, 75% of the potential pool of wind‐dispersed species has reached the polders, whereas this was only 50% for species lacking specific adaptations to long‐distance dispersal. However, the average percentage of successful colonizing species present per forest was substantially lower, ranging from 15 to 37%. Conclusions: The data strongly suggest that the colonization process in polder forests is still in its initial phase, during which easily dispersed species dominate the vegetation. Colonization success of common species that lack adaptations to long‐distance dispersal is affected by spatial configuration of the forests, and most rare species that could potentially occur in these forests are still absent. Implications for conservation of rare species in fragmented landscapes are discussed.