Integrating species traits into species pools

Despite decades of research on the species‐pool concept and the recent explosion of interest in trait‐based frameworks in ecology and biogeography, surprisingly little is known about how spatial and temporal changes in species‐pool functional diversity (SPFD) influence biodiversity and the processes underlying community assembly. Current trait‐based frameworks focus primarily on community assembly from a static regional species pool, without considering how spatial or temporal variation in SPFD alters the relative importance of deterministic and stochastic assembly processes. Likewise, species‐pool concepts primarily focus on how the number of species in the species pool influences local biodiversity. However, species pools with similar richness can vary substantially in functional‐trait diversity, which can strongly influence community assembly and biodiversity responses to environmental change. Here, we integrate recent advances in community ecology, trait‐based ecology, and biogeography to provide a more comprehensive framework that explicitly considers how variation in SPFD, among regions and within regions through time, influences the relative importance of community assembly processes and patterns of biodiversity. First, we provide a brief overview of the primary ecological and evolutionary processes that create differences in SPFD among regions and within regions through time. We then illustrate how SPFD may influence fundamental processes of local community assembly (dispersal, ecological drift, niche selection). Higher SPFD may increase the relative importance of deterministic community assembly when greater functional diversity in the species pool increases niche selection across environmental gradients. In contrast, lower SPFD may increase the relative importance of stochastic community assembly when high functional redundancy in the species pool increases the influence of dispersal history or ecological drift. Next, we outline experimental and observational approaches for testing the influence of SPFD on assembly processes and biodiversity. Finally, we highlight applications of this framework for restoration and conservation. This species‐pool functional diversity framework has the potential to advance our understanding of how local‐ and regional‐scale processes jointly influence patterns of biodiversity across biogeographic regions, changes in biodiversity within regions over time, and restoration outcomes and conservation efforts in ecosystems altered by environmental change

Local Species Diversity, Β-Diversity and Climate Influence the Regional Stability of Bird Biomass Across North America

Biodiversity often stabilizes aggregate ecosystem properties (e.g. biomass) at small spatial scales. However, the importance of species diversity within communities and variation in species composition among communities (β-diversity) for stability at larger scales remains unclear. Using a continental-scale analysis of 1657 North American breeding-bird communities spanning 20-years and 35 ecoregions, we show local species diversity and β-diversity influence two components of regional stability: local stability (stability of bird biomass within sites) and spatial asynchrony (asynchronous fluctuations in biomass among sites). We found spatial asynchrony explained three times more variation in regional stability of bird biomass than did local stability. This result contrasts with studies at smaller spatial scales—typically plant metacommunities under 1 ha—that find local stability to be more important than spatial asynchrony. Moreover, spatial asynchrony of bird biomass increased with bird β-diversity and climate heterogeneity (temperature and precipitation), while local stability increased with species diversity. Our study reveals new insights into the scale-dependent processes regulating ecosystem stability, providing evidence that both local biodiversity loss and homogenization can destabilize ecosystem processes at biogeographic scales

Negative density dependence mediates biodiversity–productivity relationships across scales

Regional species diversity generally increases with primary productivity whereas local diversity–productivity relationships are highly variable. This scale-dependence of the biodiversity–productivity relationship highlights the importance of understanding the mechanisms that govern variation in species composition among local communities, which is known as β-diversity. Hypotheses to explain changes in β-diversity with productivity invoke multiple mechanisms operating at local and regional scales, but the relative importance of these mechanisms is unknown. Here we show that changes in the strength of local density-dependent interactions within and among tree species explain changes in β-diversity across a subcontinental-productivity gradient. Stronger conspecific relative to heterospecific negative density dependence in more productive regions was associated with higher local diversity, weaker habitat partitioning (less species sorting), and homogenization of community composition among sites (lower β-diversity). Regional processes associated with changes in species pools had limited effects on β-diversity. Our study suggests that systematic shifts in the strength of local interactions within and among species might generally contribute to some of the most prominent but poorly understood gradients in global biodiversity

Beta-diversity, Environmental Change, and the Stability of Regional Ecosystems

Human activities are profoundly altering biodiversity at all spatial scales by disturbing local interaction networks, homogenizing regional biotas, and causing global extinctions. These changes in biodiversity can in turn influence the provisioning and stability of vital ecosystem functions (biomass production, biogeochemical cycling, pollination services, etc.). Therefore, a priority for ecology and global-change biology is to provide predictions for how biodiversity will respond to environmental changes. Inconveniently, biodiversity often responds unpredictably even when disturbances or environmental conditions are similar. Such contingencies have prevented the development of a general theory for ecological communities, undermining explanatory power and causing some to challenge the relevance of community ecology. This contingency may in part result from a pattern-first focus on single dimensions of biodiversity. Biodiversity is multi-dimensional, but most hypotheses to explain it have focused on the number of species (richness) or mean measures of diversity at local or regional scales (α-diversity and γ-diversity, respectively). Meanwhile, some of the largest changes in biodiversity worldwide are occurring through changes to the spatial variation in species composition (β-diversity). As a scaler that links patterns at local and regional scales, β-diversity can provide key insights into multi-scale mechanisms through which environmental change influences community assembly, biotic homogenization, and ecosystem stability. Recent conceptual frameworks suggest that four fundamental, high-level processes – speciation, dispersal, niche-selection, and ecological drift – may provide a path towards a general theory of ecological communities. However, it is unknown whether this approach will reconcile much of the contingency currently plaguing community ecology or provide useful predictions necessary to anticipate and mitigate undesired changes in biodiversity. There are two primary goals of my dissertation: 1) to understand the factors that mediate the importance of fundamental community assembly processes that cause β-diversity patterns; and 2) to determine how spatial processes that alter β-diversity contribute to, or undermine, the stability of large regional ecosystems. In Chapter 2, I address why the relative importance of selection and drift varies across natural communities for structuring herbaceous plant species and trait β-diversity. I find that drift plays an underappreciated role in causing biodiversity patterns in environmentally structured landscapes. This study highlights that understanding the scale-dependent mechanisms driving trait filtering and community size can reveal why the importance of selection and drift varies across communities and scales. In Chapter 3, I synthesize experiments to understand how and why dispersal alters the importance of drift and selection during community response to disturbance. I find that contingent assembly outcomes that cause variation in β-diversity following disturbance can be explained by dispersal that alters community size and the strength of selection. In Chapter 4, I ask how and why bird biodiversity across scales influences that stability of regional ecosystems in North America. I find that bird species β-diversity and climate heterogeneity generate asynchronous dynamics among local communities that stabilize total bird biomass at regional scales. By integrating concepts from community assembly theory, spatial ecology, and functional ecology my dissertation provides novel perspectives on the processes that influence variation in biodiversity and their consequences for ecosystem stability across scales. These insights have broad implications for both general theory and the potential to aid development of more comprehensive strategies for biodiversity conservation, ecosystem management, and landscape restoration

Functional Relationships Reveal Keystone Effects Of The Gopher Tortoise On Vertebrate Diversity In A Longleaf Pine Savanna

Keystone species are important drivers of diversity patterns in many ecosystems. Their effects on ecological processes are fundamental to understanding community dynamics, making them attractive conservation targets for ecosystem management. However, many studies assume keystone effects are constant. By developing functional relationships of species’ effects and assessing how they vary with context, we can design more efficient conservation strategies to maintain keystone impacts. The threatened gopher tortoise (Gopherus polyphemus) is presumed to be a keystone species promoting biodiversity in endangered longleaf pine ecosystems of the Southeastern Coastal Plain, USA. Although many commensals use tortoise burrows, their putative keystone influence on emergent diversity patterns lacks critical evaluation. We quantified the functional relationship between tortoise burrow density and non-volant vertebrate diversity in a longleaf pine savanna, located in central Florida. Tortoise burrow density had a positive effect on vertebrate diversity and evenness but did not affect species richness. This relationship was robust across fire disturbance regimes and was the primary factor explaining diversity at the local scale. Our results demonstrate keystone effects of the gopher tortoise through an ecosystem engineering mechanism. Continued gopher tortoise population declines will have large, negative impacts on vertebrate diversity in this biodiversity hotspot. Therefore, maintaining gopher tortoise populations is critical to effectively conserve dependent species and the function of endangered longleaf pine ecosystems. We show that developing a functional understanding of keystone relationships (not a binomial categorization) can lead to important insights into community processes

Sample Grain Influences The Functional Relationship Between Canopy Cover And Gopher Tortoise (Gopherus Polyphemus) Burrow Abandonment

Change in vegetation structure alters habitat suitability for the threatened gopher tortoise (Gopherus polyphemus). An understanding of this dynamic is crucial to inform habitat and tortoise management strategies. However, it is not known how the choice of the sample grain (i.e., cell size) at which vegetation structure is measured impacts estimates of tortoise-habitat relationships. We used lidar remote sensing to estimate canopy cover around 1573 gopher tortoise burrows at incrementally larger sample grains (1-707 m2) in 450 ha of longleaf pine (Pinus palustris) savanna. Using an information theoretic approach, we demonstrate that the choice of grain size profoundly influences modeled relationships between canopy cover and burrow abandonment. At the most supported grain size (314 m2), the probability of burrow abandonment increased by 1.7% with each percent increase in canopy cover. Ultimately, detecting the appropriate sample grain can lead to more effective development of functional relationships and improve predictive models to manage gopher tortoise habitats

Predicting wading bird and aquatic faunal responses to ecosystem restoration scenarios

In large-scale conservation decisions, scenario planning identifies key uncertainties of ecosystem function linked to ecological drivers affected by management, incorporates ecological feedbacks, and scales up to answer questions robust to alternative futures. Wetland restoration planning requires an understanding of how proposed changes in surface hydrology, water storage, and landscape connectivity affect aquatic animal composition, productivity, and food-web function. In the Florida Everglades, reintroduction of historical hydrologic patterns is expected to increase productivity of all trophic levels. Highly mobile indicator species such as wading birds integrate secondary productivity from aquatic prey (small fishes and crayfish) over the landscape. To evaluate how fish, crayfish, and wading birds may respond to alternative hydrologic restoration plans, we compared predicted small fish density, crayfish density and biomass, and wading bird occurrence for existing conditions to four restoration scenarios that varied water storage and removal of levees and canals (i.e. decompartmentalization). Densities of small fish and occurrence of wading birds are predicted to increase throughout most of the Everglades under all restoration options because of increased flows and connectivity. Full decompartmentalization goes furthest toward recreating hypothesized historical patterns of fish density by draining excess water ponded by levees and hydrating areas that are currently drier than in the past. In contrast, crayfish density declined and species composition shifted under all restoration options because of lengthened hydroperiods (i.e. time of inundation). Under full decompartmentalization, the distribution of increased prey available for wading birds shifted south, closer to historical locations of nesting activity in Everglades National Park

Integrating species traits into species pools.

Despite decades of research on the species-pool concept and the recent explosion of interest in trait-based frameworks in ecology and biogeography, surprisingly little is known about how spatial and temporal changes in species-pool functional diversity (SPFD) influence biodiversity and the processes underlying community assembly. Current trait-based frameworks focus primarily on community assembly from a static regional species pool, without considering how spatial or temporal variation in SPFD alters the relative importance of deterministic and stochastic assembly processes. Likewise, species-pool concepts primarily focus on how the number of species in the species pool influences local biodiversity. However, species pools with similar richness can vary substantially in functional-trait diversity, which can strongly influence community assembly and biodiversity responses to environmental change. Here, we integrate recent advances in community ecology, trait-based ecology, and biogeography to provide a more comprehensive framework that explicitly considers how variation in SPFD, among regions and within regions through time, influences the relative importance of community assembly processes and patterns of biodiversity. First, we provide a brief overview of the primary ecological and evolutionary processes that create differences in SPFD among regions and within regions through time. We then illustrate how SPFD may influence fundamental processes of local community assembly (dispersal, ecological drift, niche selection). Higher SPFD may increase the relative importance of deterministic community assembly when greater functional diversity in the species pool increases niche selection across environmental gradients. In contrast, lower SPFD may increase the relative importance of stochastic community assembly when high functional redundancy in the species pool increases the influence of dispersal history or ecological drift. Next, we outline experimental and observational approaches for testing the influence of SPFD on assembly processes and biodiversity. Finally, we highlight applications of this framework for restoration and conservation. This species-pool functional diversity framework has the potential to advance our understanding of how local- and regional-scale processes jointly influence patterns of biodiversity across biogeographic regions, changes in biodiversity within regions over time, and restoration outcomes and conservation efforts in ecosystems altered by environmental change