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

Changes in co-existence mechanisms along a long-term soil chronosequence revealed by functional trait diversity

1. Functional trait diversity can reveal mechanisms of species co-existence in plant communities. Few studies have tested whether functional diversity for foliar traits related to resource use strategy increases or decreases with declining soil phosphorus (P) in forest communities. 2. We quantified tree basal area and four foliar functional traits (i.e. nitrogen (N), phosphorus (P), thickness and tissue density) for all woody species along the c. 120 kyr Franz Josef soil chronosequence in cool temperate rainforest, where strong shifts occur in light and soil nutrient availability (i.e. total soil P declines from 805 to 100 mg g–1). We combined the abundance and trait data in functional diversity indices to quantify trait convergence and divergence, in an effort to determine whether mechanisms of co-existence change with soil fertility. 3. Relationships between species trait means and total soil N and P were examined using multiple regression, with and without weighting of species abundances. We used Rao’s quadratic entropy to quantify functional diversity at the plot scale, then compared this with random expectation, using a null model that randomizes abundances across species within plots. Taxonomic diversity was measured using Simpson’s Diversity. Relationships between functional and taxonomic diversity and total soil P were examined using jackknife linear regression. 4. Leaf N and P declined and leaf thickness and density increased monotonically with declining total soil P along the sequence; these relationships were unaffected by abundance-weighting of species in the analyses. Inclusion of total soil N did not improve predictions of trait means. All measures of diversity calculated from presence/absence data were unrelated to total soil N and P. There was no evidence for a relationship between Rao values using quantitative abundances and total soil P. However, there was a strongly positive relationship between Rao, expressed relative to random expectation, and total soil P, indicating trait convergence of dominant species as soil P declined. 5. Synthesis: Our results demonstrate that at high fertility dominant species differ in resource use strategy, but as soil fertility declines over the long-term, dominant species increasingly converge on a resource-retentive strategy. This suggests that differentiation in resource use strategy is required for co-existence at high fertility but not in low fertility ecosystems

Time for a plant structural economics spectrum

We argue that tree and crown structural diversity can and should be integrated in the whole-plant economics spectrum. Ecologists have found that certain functional trait combinations have been more viable than others during evolution, generating a trait trade-off continuum which can be summarized along a few axes of variation, such as the "worldwide leaf economics spectrum" and the "wood economics spectrum." However, for woody plants the crown structural diversity should be included as well in the recently introduced "global spectrum of plant form and function," which now merely focusses on plant height as structural factor. The recent revolution in terrestrial laser scanning (TLS) unlocks the possibility to describe the three dimensional structure of trees quantitatively with unprecedented detail. We demonstrate that based on TLS data, a multidimensional structural trait space can be constructed, which can be decomposed into a few descriptive axes or spectra. We conclude that the time has come to develop a "structural economics spectrum" for woody plants based on structural trait data across the globe. We make suggestions as to what structural features might lie on this spectrum and how these might help improve our understanding of tree form-function relationships

Increasing the resilience of the Australian alpine flora to climate change

 The alpine region around Australia’s highest mountain, Mt Kosciuszko, is part of one of the three most at risk ecosystems in Australia from climate change. With higher temperatures and decreased precipitation, snow cover is already declining with even greater reductions predicted in the short to medium term (2020 to 2050). Consequently the distribution of many native plants and animals may contract, while the distribution of weeds and feral animals may expand. Wildfires in the region are also likely to be more frequent and intense. To contribute to our understanding of how changes in the environment alter plant composition and ecological process, we conducted a series of functional trait analyses of existing composition datasets. We collected trait data in the field for 220 species including canopy height, leaf area, leaf dry matter content and specific leaf area (SLA). Variation in traits among the alpine flora was not related to species distributional ranges. Traits were strongly associated with growth forms, with shrubs often taller than herbs and graminoids, but often had small, tough, long-lasting leaves. Species traits were combined with relative cover values to calculate community trait weighted means, a commonly used measure of functional diversity. Functional diversity varied with altitude/duration of snow cover. For example, shrubs which are taller with small tough leaves dominated lower altitude summits, while at higher altitude summits, large, soft leaved herbs and graminoids dominated. Late lying snowpatches areas with short growing seasons were dominated by low growing herbs and graminoids with small leaves while areas with longer growing seasons were dominated by herbs and graminoids that were taller and had larger leaves. Recovery from fire differed among plant communities. The composition and functional diversity of recovering tall alpine herbfield is trending towards that of equivalent unburnt sites, while burnt windswept feldmark was colonised by graminoids and herbs that are often found in tall alpine herbfield species, with limited shrub recovery in the first nine years post fire. Grazing by feral hares had no effect on composition or functional diversity, while vegetation recovering from cattle grazing showed clear changes in composition and functional diversity even 43 years later. Prioritising management for this high value conservation region, therefore, involves enhancing resilience by minimising existing threats, particularly those from fire, weeds and hard-hooved grazing animals which will be exacerbate by climate change.Please cite as: Pickering, C, & Venn, S, 2013 Increasing the resilience of the Australian flora to climate change and associated threats: a plant functional traits approach National Climate Change Adaptation Research Facility, Gold Coast, pp. 94 Abstract The alpine region around Australia’s highest mountain, Mt Kosciuszko, is part of one of the three most at risk ecosystems in Australia from climate change. With higher temperatures and decreased precipitation, snow cover is already declining with even greater reductions predicted in the short to medium term (2020 to 2050). Consequently the distribution of many native plants and animals may contract, while the distribution of weeds and feral animals may expand. Wildfires in the region are also likely to be more frequent and intense. To contribute to our understanding of how changes in the environment alter plant composition and ecological process, we conducted a series of functional trait analyses of existing composition datasets. We collected trait data in the field for 220 species including canopy height, leaf area, leaf dry matter content and specific leaf area (SLA). Variation in traits among the alpine flora was not related to species distributional ranges. Traits were strongly associated with growth forms, with shrubs often taller than herbs and graminoids, but often had small, tough, long-lasting leaves. Species traits were combined with relative cover values to calculate community trait weighted means, a commonly used measure of functional diversity. Functional diversity varied with altitude/duration of snow cover. For example, shrubs which are taller with small tough leaves dominated lower altitude summits, while at higher altitude summits, large, soft leaved herbs and graminoids dominated. Late lying snowpatches areas with short growing seasons were dominated by low growing herbs and graminoids with small leaves while areas with longer growing seasons were dominated by herbs and graminoids that were taller and had larger leaves. Recovery from fire differed among plant communities. The composition and functional diversity of recovering tall alpine herbfield is trending towards that of equivalent unburnt sites, while burnt windswept feldmark was colonised by graminoids and herbs that are often found in tall alpine herbfield species, with limited shrub recovery in the first nine years post fire. Grazing by feral hares had no effect on composition or functional diversity, while vegetation recovering from cattle grazing showed clear changes in composition and functional diversity even 43 years later. Prioritising management for this high value conservation region, therefore, involves enhancing resilience by minimising existing threats, particularly those from fire, weeds and hard-hooved grazing animals which will be exacerbate by climate change

Functional biogeography of oceanic islands and the scaling of functional diversity in the Azores

Analyses of species-diversity patterns of remote islands have been crucial to the development of biogeographic theory, yet little is known about corresponding patterns in functional traits on islands and how, for example, they may be affected by the introduction of exotic species. We collated trait data for spiders and beetles and used a functional diversity index (FRic) to test for nonrandomness in the contribution of endemic, other native (also combined as indigenous), and exotic species to functional-trait space across the nine islands of the Azores. In general, for both taxa and for each distributional category, functional diversity increases with species richness, which, in turn scales with island area. Null simulations support the hypothesis that each distributional group contributes to functional diversity in proportion to their species richness. Exotic spiders have added novel trait space to a greater degree than have exotic beetles, likely indicating greater impact of the reduction of immigration filters and/or differential historical losses of indigenous species. Analyses of species occurring in native-forest remnants provide limited indications of the operation of habitat filtering of exotics for three islands, but only for beetles. Although the general linear (not saturating) pattern of trait-space increase with richness of exotics suggests an ongoing process of functional enrichment and accommodation, further work is urgently needed to determine how estimates of extinction debt of indigenous species should be adjusted in the light of these findings

Using a Multi-Trait Approach to Manipulate Plant Functional Diversity in a Biodiversity-Ecosystem Function Experiment

A frequent pattern emerging from biodiversity-ecosystem function studies is that functional group richness enhances ecosystem functions such as primary productivity. However, the manipulation of functional group richness goes along with major disadvantages like the transformation of functional trait data into categories or the exclusion of functional differences between organisms in the same group. In a mesocosm study we manipulated plant functional diversity based on the multi-trait Functional Diversity (FD)-approach of Petchey and Gaston by using database data of seven functional traits and information on the origin of the species in terms of being native or exotic. Along a gradient ranging from low to high FD we planted 40 randomly selected eight-species mixtures under controlled conditions. We found a significant positive linear correlation of FD with aboveground productivity and a negative correlation with invasibility of the plant communities. Based on community-weighted mean calculations for each functional trait, we figured out that the traits N-fixation and species origin, i.e. being native or exotic, played the most important role for community productivity. Our results suggest that the identification of the impact of functional trait diversity and the relative contributions of relevant traits is essential for a mechanistic understanding of the role of biodiversity for ecosystem functions such as aboveground biomass production and resistance against invasion

Incorporating intraspecific trait variation into functional diversity: Impacts of selective logging on birds in Borneo

1. As conservation increasingly recognises the importance of species’ functional roles in ecosystem processes, studies are shifting away from measuring species richness towards measures that account for the functional differences between species in a community. These functional diversity (FD) indices have received much recent attention and refinement, but their greatest limitation remains their inability to incorporate information about intraspecific trait variation (ITV). 2. We use an individual-based model to account for ITV when calculating the functional diversity of two avian communities in Borneo; one in primary (unlogged) forest and one in selectively logged forest. We deal with the scarcity of trait data for individual species by developing a simulation approach, taking data from the literature where necessary. Using a bootstrapping procedure, we produce a range of ecologically feasible FD values taking account of ITV for five commonly-used FD indices, and we quantify the confidence that can be placed in these values using a newly-developed bootstrapping method: btFD. 3. We found that incorporating ITV significantly altered the FD values of all indices used in our models. The rank order of FD for the two communities, indicating whether diversity was higher in primary or selectively logged forest, was largely unchanged by the inclusion of ITV. However, by accounting for ITV, we were able to reveal previously unrecognized impacts of selective logging on avian functional diversity through a narrower dispersion of individuals in functional trait space in logged forest. 4. Our results highlight the importance of incorporating ITV into measures of functional diversity, whilst our simulation approach addresses the frequently encountered difficulty of working with sparse trait data and quantifies the confidence that should be placed in such findings

The road to functional recovery : temporal effects of matrix regeneration on Amazonian bats

Across the tropics, vast deforested areas are undergoing forest regeneration due to land abandonment. Although secondary forest is an expanding type of landscape matrix that has been shown to buffer some of the negative consequences of forest loss and fragmentation on taxonomic diversity, little is known in this regard about the functional dimension of biodiversity. We took advantage of an ecosystem-wide fragmentation experiment to investigate longer term changes in functional diversity of a mega-diverse Amazonian bat assemblage associated with regrowth development in the matrix. We found that matrix regeneration affected several facets of bat functional diversity in secondary forest over time, increasing functional α diversity, species- and community-level functional uniqueness, altering functional trait composition, and resulting in functional β-diversity changes via trait gains. However, approximately 30 years of matrix regeneration were insufficient for functional diversity to recover to the same levels as in continuous forest. Our results suggest that a combination of natural, human-assisted, and active restoration is likely to be the most successful strategy for restoring functional biodiversity of bats in human-modified tropical landscapes, a finding that most likely also applies to many other taxa

Elevation, moisture and shade drive the functional and phylogenetic meadow communities’ assembly in the northeastern Tibetan Plateau

Despite a long history of alpine meadows studies, uncertainty remains about the importance of environmental factors in structuring their assembly. We examined the functional and phylogenetic structure of 170 alpine Tibetan meadow communities in relation to elevation, soil moisture and shade. Functional community structure was estimated with both communityweighted mean (CWM) trait values for specific leaf area (SLA), plant height and seed mass and functional diversity (Rao’s quadratic index) for their traits individually and in combination (multivariate functional diversity). We found that shade induced by woody plants significantly increased the phylogenetic diversity and functional diversity of SLA of co-occurring species, suggesting that woody plants behave as “ecosystem engineers” creating a different environment that allows the existence of shade tolerant species and thereby facilitates the coexistence of plant species with different light resource acquisition strategies. We also found evidence for a clear decrease in phylogenetic diversity, CWM and functional diversity related to plant height in the two extreme, both the dry and wet, soil moisture conditions. This indicates that both drought and excess moisture may act as environmental filters selecting species with close phylogenetic relationships and similar height. Moreover, we detected significant decreases in both CWM and functional diversity for seed mass along elevational gradients, suggesting that low net primary productivity (NPP) limits seed size. Finally, because of different individual trait responses to environmental factors, the multivariate functional diversity did not change across environmental gradients. This lack of multivariate response supports the hypothesis that multiple processes, such as environmental filtering, competition and facilitation, may operate simultaneously and exert opposing effects on community assembly along different niche (e.g., water use, light acquisition) axes, resulting in no overall functional community structure change. This contrast between individual and multivariate trait patterns highlights the importance of examining individual traits linked with different ecological processes to better understand the mechanisms of community assembly