Dendritic network structure and dispersal affect temporal dynamics of diversity and species persistence

Landscape connectivity structure, specifically the dendritic network structure of rivers, is expected to influence community diversity dynamics by altering dispersal patterns, and subsequently the unfolding of species interactions. However, previous comparative and experimental work on dendritic metacommunities has studied diversity mostly from an equilibrium perspective. Here we investigated the effect of dendritic versus linear network structure on local (α‐diversity), among (β‐diversity) and total (γ‐diversity) temporal species community diversity dynamics. Using a combination of microcosm experiments, which allowed for active dispersal of 14 protists and a rotifer species, and numerical analyses, we demonstrate the general importance of spatial network configuration and basic life history tradeoffs as driving factors of different diversity patterns in linear and dendritic systems. We experimentally found that community diversity patterns were shaped by the interaction of dispersal within the networks and local species interactions. Specifically, α‐diversity remained higher in dendritic networks over time, especially at highly connected sites. β‐diversity was initially greater in linear networks, due to increased dispersal limitation, but became more similar to β‐diversity in dendritic networks over time. Comparing the experimental results with a neutral metacommunity model we found that dispersal and network connectivity alone may, to a large extent, explain α‐ and β‐diversity dynamics. However, additional mechanisms, such as variation in carrying capacity and competition–colonization tradeoffs, were needed in the model to capture the detailed temporal diversity dynamics of the experiments, such as a general decline in γ‐diversity and long‐term dynamics in α‐diversity.     

Diversity in riverine metacommunities: a network perspective

The influence of spatial processes on diversity and community dynamics is generally recognized in ecology and also applied to conservation projects involving forest and grassland ecosystems. Riverine ecosystems, however, have been for a long time viewed from a local or linear perspective, even though the treelike branching of river networks is universal. River networks (so-called dendritic networks) are not only structured in a hierarchic way, but the dendritic landscape structure and physical flows often dictate distance and directionality of dispersal. Theoretical models suggest that the specific riverine network structure directly affects diversity patterns. Recent experimental and comparative data are supporting this idea. Here, I provide an introduction on theoretical findings suggesting that genetic diversity, heterozygosity and species richness are higher in dendritic systems compared to linear or two-dimensional lattice landscapes. The characteristic diversity patterns can be explained in a network perspective, which also offers universal metrics to better understand and protect riverine diversity. I show how appropriate metrics describing network centrality and dispersal distances are superior to classic measures still applied in aquatic ecology, such as Strahler order or Euclidian distance. Finally, knowledge gaps and future directions of research are identified. The network perspective employed here may help to generalize findings on riverine biodiversity research and can be applied to conservation and river restoration projects.     

Classical metapopulation dynamics and eco‐evolutionary feedbacks in dendritic networks

Eco‐evolutionary dynamics are now recognized to be highly relevant for population and community dynamics. However, the impact of evolutionary dynamics on spatial patterns, such as the occurrence of classical metapopulation dynamics, is less well appreciated. Here, we analyse the evolutionary consequences of spatial network connectivity and topology for dispersal strategies and quantify the eco‐evolutionary feedback in terms of altered classical metapopulation dynamics. We find that network properties, such as topology and connectivity, lead to predictable spatio‐temporal correlations in fitness expectations. These spatio‐temporally stable fitness patterns heavily impact evolutionarily stable dispersal strategies and lead to eco‐evolutionary feedbacks on landscape level metrics, such as the number of occupied patches, the number of extinctions and recolonizations as well as metapopulation extinction risk and genetic structure. Our model predicts that classical metapopulation dynamics are more likely to occur in dendritic networks, and especially in riverine systems, compared to other types of landscape configurations. As it remains debated whether classical metapopulation dynamics are likely to occur in nature at all, our work provides an important conceptual advance for understanding the occurrence of classical metapopulation dynamics which has implications for conservation and management of spatially structured populations.     

Importance of scale,land‐use,and stream network properties for riparian plant communities along an urban gradient

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Effects of colonization asymmetries on metapopulation persistence

Ocean currents, prevailing winds, and the hierarchical structures of river networks are known to create asymmetries in re-colonization between habitat patches. The impacts of such asymmetries on metapopulation persistence are seldom considered, especially rarely in theoretical studies. Considering three classical models (the island, the stepping stone and the distance-dependent model), we explore how metapopulation persistence is affected by (i) asymmetry in dispersal strength, in which the colonization rate between two patches differs in direction, and (ii) asymmetry in connectivity, in which the overall colonization pattern displays asymmetry (circulating or dendritic networks). Viability can be drastically reduced when directional bias in dispersal strength is higher than 25%. Re-colonization patterns that allow for strong local connectivity provide the highest persistence compared to systems that allow circulation. Finally, asymmetry has relatively weak effects when metapopulations maintain strong general connectivity.     

Contrasting patterns and mechanisms of spatial turnover for native and exotic freshwater fish in Europe

Aim We compare the distribution patterns of native and exotic freshwater fish in Europe, and test whether the same mechanisms (environmental filtering and/or dispersal limitation) govern patterns of decrease in similarity of native and exotic species composition over geographical distance (spatial species turnover). Locations Major river basins of Europe. Methods Data related to geography, habitat diversity, regional climate and species composition of native and exotic freshwater fish were collated for 26 major European river basins. We explored the degree of nestedness in native and exotic species composition, and quantified compositional similarity between river basins according to the beta‐sim (independent of richness gradient) and Jaccard (dependent of richness gradient) indices of similarity. Multiple regression on distance matrices and variation‐partitioning approaches were used to quantify the relative roles of environmental filtering and dispersal limitation in shaping patterns of decreasing compositional similarity over geographical distance. Results Native and exotic species exhibited significant nested patterns of species composition, indicating that differences in fish species composition between river basins are primarily the result of species loss, rather than species replacement. Both native and exotic compositional similarity decreased significantly with increasing geographical distance between river basins. However, gradual changes in species composition with geographical distance were found only for exotic species. In addition, exotic species displayed a higher rate of similarity decay (higher species turnover rate) with geographical distance, compared with native species. Lastly, the majority of explained variation in exotic compositional similarity was uniquely related to geography, whereas native compositional similarity was either uniquely explained by geography or jointly explained by environment and geography. Main conclusions Our study suggests that large‐scale patterns of spatial turnover for exotic freshwater fish in Europe are generated by human‐mediated dispersal limitation, whereas patterns of spatial turnover for native fish result from both dispersal limitation relative to historical events (isolation by mountain ranges, glacial history) and environmental filtering.     

Dispersal,diversity and distribution patterns in pioneer vegetation: The role of river‐floodplain connectivity

Question: Are there changes in dispersal patterns in floodplain pioneer vegetation with effects on seedling number, species richness and species composition along a gradient of declining river‐floodplain connectivity? Location: Middle Elbe river floodplain, Germany. Methods: An experiment with five treatments was set up along a gradient of declining river‐floodplain connectivity, partitioning seedlings into three groups: (1) emerging solely from water dispersed seeds, (2) from wind/animal dispersed seeds and (3) from the soil diaspore bank. Two controls were established: without any manipulation and exclusion of all seeds. The results were compared with those of vegetation and soil sampling to evaluate the representativeness of the experimental sites in terms of species composition, diversity, seedling number and soil parameters. Results: Water dispersal and the soil diaspore bank were the major dispersal strategies shaping floodplain pioneer vegetation at the Middle Elbe river. The number of seedlings, species richness and the variation in species composition in these habitats depend on the degree of connectivity. The seedling number and species richness is highest in sites of permanent or almost permanent exchange with the main channel, where water dispersal additionally contributes to the number of seedlings grown from the soil seed bank. Conclusion: The results underline the importance of river‐floodplain ecotones as sink habitats for water‐dispersed seeds. Considering the strongly reduced river‐floodplain interactions due to dykes and other engineering structures, management strategies are necessary to improve connectivity and the renewal of fluvial land forms.     

Spatial eigenfunction analyses in stream networks: do watercourse and overland distances produce different results?

1. The use of spatial variables is a common procedure in ecological studies. The technique is based on the definition of a connectivity/distance matrix that conceptually defines the dispersal of organisms. The shortest distance between two points is a straight line. Despite the fact that a straight line may not represent the easiest dispersal path for many kinds of organisms, straight‐line distances are often used to detect patterns. We argue that other types of connectivity/distance matrices will better represent dispersal paths, such as the watercourse distance for aquatic organisms (e.g. fish, shrimps). 2. We used empirical and simulated community data to evaluate the usefulness of spatial variables generated from watercourse and overland (straight‐line) distances. 3. Spatial variables based on watercourse distances captured patterns that straight‐line distances did not, and provided better representations of the spatial patterns generated by dispersal along a dendritic network.     

The role of environmental and spatial processes in structuring native and non‐native fish communities across thousands of lakes

Quantifying the role of spatial patterns is an important goal in ecology to further understand patterns of community composition. We quantified the relative role of environmental conditions and regional spatial patterns that could be produced by environmental filtering and dispersal limitation on fish community composition for thousands of lakes. A database was assembled on fish community composition, lake morphology, water quality, climatic conditions, and hydrological connectivity for 9885 lakes in Ontario, Canada. We utilized a variation partitioning approach in conjunction with Moran's Eigenvector Maps (MEM) and Asymmetric Eigenvector Maps (AEM) to model spatial patterns that could be produced by human‐mediated and natural modes of dispersal. Across 9885 lakes and 100 fish species, environmental factors and spatial structure explained approximately 19% of the variation in fish community composition. Examining the proportional role of spatial structure and environmental conditions revealed that as much as 90% of the explained variation in native species assemblage composition is governed by environmental conditions. Conversely on average, 67% of the explained variation in non‐native assemblage composition can be related to human‐mediated dispersal. This study highlights the importance of including spatial structure and environmental conditions when explaining patterns of community composition to better discriminate between the ecological processes that underlie biogeographical patterns of communities composed of native and non‐native fish species.     

The importance of landscape and habitat properties in explaining instantaneous and long‐term distributions of large branchiopods in subtropical temporary pans

1. The notion that the spatial configuration of habitat patches has to be taken into account to understand the structure and dynamics of ecological communities is the starting point of metacommunity ecology. One way to assess metacommunity structure is to investigate the relative importance of environmental heterogeneity and spatial structure in explaining community patterns over different spatial and temporal scales. 2. We studied metacommunity structure of large branchiopod assemblages characteristic of subtropical temporary pans in SE Zimbabwe using two community data sets: a community snapshot and a long‐term data set covering 4 years. We assessed the relative importance of environmental heterogeneity and dispersal (inferred from patch occupancy patterns) as drivers of community structure. Furthermore, we contrasted metacommunity patterns in pans that occasionally connect to the river (floodplain pans) and pans that lack such connections altogether (endorheic pans) using redundancy models. 3. Echoes of species sorting and dispersal limitation emerge from our data set, suggesting that both local and regional processes contribute to explaining branchiopod assemblages in this system. Relative importance of local and regional factors depended on the type of data set considered. Overall, habitat characteristics that vary in time, such as conductivity, hydroperiod and vegetation cover, best explained the instantaneous species composition observed during a snapshot sampling while long‐term species composition appeared to be linked to more constant intrinsic habitat properties such as river connectivity and spatial location.     

Seeing through the static: the temporal dimension of plant–animal mutualistic interactions

Most studies of plant–animal mutualistic networks have come from a temporally static perspective. This approach has revealed general patterns in network structure, but limits our ability to understand the ecological and evolutionary processes that shape these networks and to predict the consequences of natural and human‐driven disturbance on species interactions. We review the growing literature on temporal dynamics of plant–animal mutualistic networks including pollination, seed dispersal and ant defence mutualisms. We then discuss potential mechanisms underlying such variation in interactions, ranging from behavioural and physiological processes at the finest temporal scales to ecological and evolutionary processes at the broadest. We find that at the finest temporal scales (days, weeks, months) mutualistic interactions are highly dynamic, with considerable variation in network structure. At intermediate scales (years, decades), networks still exhibit high levels of temporal variation, but such variation appears to influence network properties only weakly. At the broadest temporal scales (many decades, centuries and beyond), continued shifts in interactions appear to reshape network structure, leading to dramatic community changes, including loss of species and function. Our review highlights the importance of considering the temporal dimension for understanding the ecology and evolution of complex webs of mutualistic interactions.     

Contrasting metacommunity structure and beta diversity in an aquatic‐floodplain system

Habitat connectivity and dispersal interact to structure metacommunities, but few studies have examined these patterns jointly for organisms across the aquatic–terrestrial ecotone. We assessed metacommunity structure and beta diversity patterns of instream benthic invertebrates, riparian carabid beetles (Order: Coleoptera; Family: Carabidae) and riparian spiders (Order: Araneae) at fifteen sites in a river‐floodplain system. Sampling took place over a three‐year period (2010–2012) in the Rhine‐Main‐Observatory LTER site on the Kinzig River, central Germany. This allowed disentangling the combined influence, and temporal variability, of habitat connectivity (i.e. between aquatic and terrestrial) and dispersal ability (i.e. between spiders and beetles, and aerial and aquatic dispersing invertebrates) on the dominant paradigms structuring these metacommunities. We found mostly consistent differences in the manner that metacommunities were structured between groups, with lower levels of variability explained for beetles compared to the other groups. Beetles were consistently structured more by turnover than nestedness components, with greater beta diversity than expected by chance and a minor spatial compared to environmental signal emerging with variance partitioning. Conversely, spiders and benthic invertebrates had lower beta diversity and greater nestedness than null expectation, and a clearer spatial signal controlling metacommunity structure. Our results suggest varying levels of mass effects and species sorting shape river‐floodplain metacommunities, depending on habitat connectivity and dispersal ability. That is, greater connectivity and lower fragmentation along the river compared to the terrestrial zone promoted mass effects, and differences in overall dispersal ability and mode (i.e. active and passive) for instream and riparian communities shifted paradigms between mass effects and species sorting.     

Production of diaspores at the landscape level regulates local colonization: an experiment with a spore‐dispersed moss

Effective dispersal is crucial to species inhabiting transient substrates in order for them to be able to persist in a landscape. Bryophytes, pteridophytes, lichens and fungi all have wind‐dispersed small diaspores and can be efficiently dispersed if their diaspores reach air masses above canopy height. However, empirical data on dispersal over landscape scales are scarce. We investigated how the colonization of an acrocarpous clay‐inhabiting pioneer moss, Discelium nudum, varied between sites that differed in connectivity to potential dispersal sources at spatial scales from 1 to 20 km in a region in northern Sweden. We recorded the colonization on ?25 introduced clay heaps at each of 14 experimental sites some months after the dispersal period. The colonization rate ranged from 0–82% and had a statistically significant relationship with a proxy for potential habitats (amount of clay‐dominated soil) in a buffer of 20 km radius surrounding the experimental sites (and also weakly with the amount of substrate in a 10 km buffer). There were no significant relationships between colonization rate and connectivity at smaller scales (1 and 5 km). We made a rough estimate of the number of spores available for dispersal in a landscape, given the amount of clay‐dominated soil, by recording the number of Discelium nudum colonies in two 25 × 25 km landscapes. The estimated available spore numbers in the different 20 km buffers were of the same order of magnitude as the deposition densities at the experimental sites calculated from the colonization rates. The results suggest that the spores of species with scattered occurrences and small diaspores (25 μm) in open landscapes can be deposited over extensive areas, at rates high enough to drive colonization patterns. This also implies that regional connectivity may be more important than local connectivity for these kinds of species.     

Proximity of restored hedgerows interacts with local floral diversity and species' traits to shape long‐term pollinator metacommunity dynamics

Disconnected habitat fragments are poor at supporting population and community persistence; restoration ecologists, therefore, advocate for the establishment of habitat networks across landscapes. Few empirical studies, however, have considered how networks of restored habitat patches affect metacommunity dynamics. Here, using a 10‐year study on restored hedgerows and unrestored field margins within an intensive agricultural landscape, we integrate occupancy modelling with network theory to examine the interaction between local and landscape characteristics, habitat selection and dispersal in shaping pollinator metacommunity dynamics. We show that surrounding hedgerows and remnant habitat patches interact with the local floral diversity, bee diet breadth and bee body size to influence site occupancy, via colonisation and persistence dynamics. Florally diverse sites and generalist, small‐bodied species are most important for maintaining metacommunity connectivity. By providing the first in‐depth assessment of how a network of restored habitat influences long‐term population dynamics, we confirm the conservation benefit of hedgerows for pollinator populations and demonstrate the importance of restoring and maintaining habitat networks within an inhospitable matrix.     

Linking dendritic network structures to population demogenetics: The downside of connectivity

Spatial structures strongly influence ecological processes. Connectivity is known to positively influence metapopulation demography and genetics by increasing the rescue effect and thus favoring individual and gene flow between populations. This result has not been tested in the special case of dendritic networks, which encompass stream and cave ecosystem for instance. We propose a first approach using an individual based model to explore the population demography and genetics in various dendritic networks. To do so, we first generate a large number of different networks, and we analyze the relationship between their hydrographical characteristics and connectivity. We show that connectivity mean and variance of connectivity are strongly correlated in dendritic networks. Connectivity segregates two types of networks: Hortonian and non‐Hortonian networks. We then simulate the population dynamics for a simple life cycle in each of the generated networks, and we analyze persistence time as well as populations structure at quasi‐stationary state. Our main results show that connectivity in dendritic networks can promote local extinction and genetic isolation by distance at low dispersal and diminish the size of the metapopulation at high dispersal. We discuss these unexpected findings in the light of connectivity spatial distribution in dendritic networks in the case of our model.     

Incorporating seed fate into plant–frugivore networks increases interaction diversity across plant regeneration stages

Plant–animal mutualistic interactions, such as pollination and seed dispersal, affect ecosystem functioning by driving plant population dynamics. However, little is known of how the diversity of interactions in these mutualistic networks determines plant regeneration dynamics. To fill this gap, interaction networks should not only account for the number of seeds dispersed by animals, but also for seed fate after dispersal. Here, we compare plant–animal networks at both the seed dispersal and seedling recruitment stage to evaluate how interaction diversity, represented by different network metrics, changes throughout the process of plant regeneration. We focused on a system with six species of frugivorous birds and three species of fleshy‐fruited trees in the temperate secondary forest of the Cantabrian Range (northern Iberian Peninsula). We considered two plant cohorts corresponding to two fruiting years showing strong differences in fruit and frugivore abundance. Seed dispersal interactions were estimated from a spatially‐explicit, field‐validated model predicting tree and bird species‐specific seed deposition in different microhabitats. These interactions were further transformed into interactions at the seedling recruitment stage by accounting for plant‐ and microhabitat‐specific seed fates estimated from field sampling. We found that network interaction diversity varied across plant regeneration stages and cohorts, both in terms of the evenness and the number of paired interactions. Tree–bird interactions were more evenly distributed across species pairs at the recruitment stage than at the seed deposition stage, although some interactions disappeared in the seed‐to‐seedling transition for one plant cohort. The variations in interaction diversity were explained by between‐plant differences in post‐dispersal seed fate and in inter‐annual fruit production, rather than by differences between frugivores in seed deposition patterns. These results highlight the need for integrating plant traits and disperser quality to predict the functional outcome of plant–animal mutualistic networks.     

A potential role for overland dispersal in shaping aquatic invertebrate communities in arid regions

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A neutral metapopulation model of biodiversity in river networks

In this paper, we develop a stochastic, discrete, structured metapopulation model to explore the dynamics and patterns of biodiversity of riparian vegetation. In the model, individual plants spread along a branched network via directional dispersal and undergo neutral ecological drift. Simulation results suggest that in comparison to 2-D landscapes with non-directional dispersal, river networks with directional dispersal have lower local (alpha) and overall (gamma) diversities, but higher between-community (beta) diversity, implying that riparian species are distributed in a more localized pattern and more vulnerable to local extinction. The relative abundance patterns also change, such that higher percentages of species are in low-abundance, or rare, classes, accompanied by concave rank-abundance curves. In contrast to existing theories, the results suggest that in river networks, increased directional dispersal reduces alpha diversity. These altered patterns and trends result from the combined effects of directionality of dispersal and river network structure, whose relative importance is in need of continuing study. In addition, riparian communities obeying neutral dynamics seem to exhibit abrupt changes where large tributaries confluence; this pattern may provide a signature to identify types of interspecific dynamics in river networks.     

Effects of branching spatial structure and life history on the asymptotic growth rate of a population

The dendritic structure of a river network creates directional dispersal and a hierarchical arrangement of habitats. These two features have important consequences for the ecological dynamics of species living within the network. We apply matrix population models to a stage-structured population in a network of habitat patches connected in a dendritic arrangement. By considering a range of life histories and dispersal patterns, both constant in time and seasonal, we illustrate how spatial structure, directional dispersal, survival, and reproduction interact to determine population growth rate and distribution. We investigate the sensitivity of the asymptotic growth rate to the demographic parameters of the model, the system size, and the connections between the patches. Although some general patterns emerge, we find that a species’ modes of reproduction and dispersal are quite important in its response to changes in its life history parameters or in the spatial structure. The framework we use here can be customized to incorporate a wide range of demographic and dispersal scenarios.     

Habitat connectivity and dispersal ability drive the assembly mechanisms of macroinvertebrate communities in river networks

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