Biodiversity in changing environments: An external-driver internal-topology framework to guide intervention.

Accompanying the climate crisis is the more enigmatic biodiversity crisis. Rapid reorganization of biodiversity due to global environmental change has defied prediction and tested the basic tenets of conservation and restoration. Conceptual and practical innovation is needed to support decision making in the face of these unprecedented shifts. Critical questions include: How can we generalize biodiversity change at the community level? When are systems able to reorganize and maintain integrity, and when does abiotic change result in collapse or restructuring? How does this understanding provide a template to guide when and how to intervene in conservation and restoration? To this end, we frame changes in community organization as the modulation of external abiotic drivers on the internal topology of species interactions, using plant-plant interactions in terrestrial communities as a starting point. We then explore how this framing can help translate available data on species abundance and trait distributions to corresponding decisions in management. Given the expectation that community response and reorganization are highly complex, the external-driver internal-topology (EDIT) framework offers a way to capture general patterns of biodiversity that can help guide resilience and adaptation in changing environments.

Seed bank bias: Differential tracking of functional traits in the seed bank and vegetation across a gradient.

A goal in trait-based ecology is to understand and predict plant community responses to environmental change; however, diversity stored within seed banks that may expand or limit these responses is typically overlooked. If seed banks store attributes that are more advantageous or vulnerable under future conditions, they could impact community adaptability to change and disturbance. We explored compositional differences between seed banks and vegetation (i.e., seed bank bias) across a 12-site gradient of increasingly higher and older soil terraces, asking: How do seed banks contribute to taxonomic and functional composition, and what do shifts in seed bank biases along the gradient (i.e., tracking) reveal about the processes driving seed bank variation and its implications for community adaptability? Across the gradient, seed banks stored distinct pools of species that added to species richness but not functional dispersion. Seed banks were generally biased toward short-life histories and "fast" species with small seeds, thinner and more acquisitive roots, and lower root biomass allocation; however, trait means in the seed bank and vegetation sometimes shifted along the gradient, amplifying or reversing these biases. For example, species with higher specific leaf area (tied to rapid resource acquisition) tended to dominate vegetation on lower soil terraces, but were more common in the seed bank on higher terraces-at least when patterns were weighted by species' relative abundances. Although seed banks were generally characterized by "fast" attributes, observed shifts in seed bank biases across the gradient-particularly in leaf traits-demonstrate that environment can impact stored diversity and, consequently, our expectations for future vegetative turnover. The seed bank bias patterns that we characterized could be the result of many potential processes, including environment- or trait-driven variation in seed bank inputs (seed production, dispersal) or losses (seed desiccation, germination), and may have important implications for a system's adaptive capacity. Only by integrating seed banks into the functional ecology agenda will we be able to unpack these processes and use seed banks more effectively in both prediction and ecosystem management.

Plant traits are differentially linked to performance in a semiarid ecosystem.

A central principle in trait-based ecology is that trait variation has an adaptive value. However, uncertainty over which plant traits influence individual performance across environmental gradients may limit our ability to use traits to infer ecological processes at larger scales. To better understand which traits are linked to performance under different precipitation regimes, we measured above- and belowground traits, growth, and reproductive allocation for four annual and four perennial species from a coastal sage scrub community in California under conditions of 50%, 100%, and 150% ambient precipitation. Across water treatments, annual species displayed morphological trait values consistent with high rates of resource acquisition (e.g., low leaf mass per area, low root tissue density, high specific root length), and aboveground measures of resource acquisition (including photosynthetic rate and leaf N concentration) were positively associated with plant performance (reproductive allocation). Results from a structural equation model demonstrated that leaf traits explained 38% of the variation in reproductive allocation across the water gradient in annual species, while root traits accounted for only 6%. Although roots play a critical role in water uptake, more work is needed to understand the mechanisms by which root trait variation can influence performance in water-limited environments. Perennial species showed lower trait plasticity than annuals across the water gradient and were more variable as a group in terms of trait-performance relationships, indicating that species rely on different functional strategies to respond to drought. Our finding that species identity drives much of the variation in trait values and trait-performance relationships across a water gradient may simplify efforts to model ecological processes, such as productivity, that are potentially influenced by environmentally induced shifts in trait values.

Leaf traits and performance vary with plant age and water availability in Artemisia californica.

BACKGROUND AND AIMS: Leaf functional traits are strongly tied to growth strategies and ecological processes across species, but few efforts have linked intraspecific trait variation to performance across ontogenetic and environmental gradients. Plants are believed to shift towards more resource-conservative traits in stressful environments and as they age. However, uncertainty as to how intraspecific trait variation aligns with plant age and performance in the context of environmental variation may limit our ability to use traits to infer ecological processes at larger scales. METHODS: We measured leaf physiological and morphological traits, canopy volume and flowering effort for Artemisia californica (California sagebrush), a dominant shrub species in the coastal sage scrub community, under conditions of 50, 100 and 150 % ambient precipitation for 3 years. KEY RESULTS: Plant age was a stronger driver of variation in traits and performance than water availability. Older plants demonstrated trait values consistent with a more conservative resource-use strategy, and trait values were less sensitive to drought. Several trait correlations were consistent across years and treatments; for example, plants with high photosynthetic rates tended to have high stomatal conductance, leaf nitrogen concentration and light-use efficiency. However, the trade-off between leaf construction and leaf nitrogen evident in older plants was absent for first-year plants. While few traits correlated with plant growth and flowering effort, we observed a positive correlation between leaf mass per area and performance in some groups of older plants. CONCLUSIONS: Overall, our results suggest that trait sensitivity to the environment is most visible during earlier stages of development, after which intraspecific trait variation and relationships may stabilize. While plant age plays a major role in intraspecific trait variation and sensitivity (and thus trait-based inferences), the direct influence of environment on growth and fecundity is just as critical to predicting plant performance in a changing environment.

Rapid root responses of seedlings exposed to a postdrought water pulse.

PREMISE OF THE STUDY: Mediterranean-type climate ecosystems experience significant variability in precipitation within and across years and may be characterized by periods of extreme drought followed by a brief, high-intensity precipitation pulse. Rapid root growth could be a key factor in effective utilization of precipitation pulses, leading to higher rates of seedling establishment. Changes in root growth rate are rarely studied, however, and patterns in seedling root traits are not well explored. We investigated the influence of an extreme postdrought precipitation event on seedlings that occur in southern California coastal sage scrub. METHODS: We measured root elongation rate, root tip appearance rate, new leaf appearance rate, and canopy growth rate on 18 mediterranean species from three growth forms. KEY RESULTS: Root elongation rate responded more strongly to the precipitation pulse than did root tip appearance rate and either metric of aboveground growth. The majority of species exhibited a significant change in root growth rate within 1 week of the pulse. Responses varied in rapidity and magnitude across species, however, and were not generally predictable based on growth form. CONCLUSIONS: While the majority of species exhibited shifts in belowground growth following the pulse, the direction and magnitude of these morphological responses were highly variable within growth form. Understanding the implications of these different response strategies for plant fitness is a crucial next step to forecasting community dynamics within ecosystems characterized by resource pulses.

Revisiting the Holy Grail: using plant functional traits to understand ecological processes.

One of ecology's grand challenges is developing general rules to explain and predict highly complex systems. Understanding and predicting ecological processes from species' traits has been considered a 'Holy Grail' in ecology. Plant functional traits are increasingly being used to develop mechanistic models that can predict how ecological communities will respond to abiotic and biotic perturbations and how species will affect ecosystem function and services in a rapidly changing world; however, significant challenges remain. In this review, we highlight recent work and outstanding questions in three areas: (i) selecting relevant traits; (ii) describing intraspecific trait variation and incorporating this variation into models; and (iii) scaling trait data to community- and ecosystem-level processes. Over the past decade, there have been significant advances in the characterization of plant strategies based on traits and trait relationships, and the integration of traits into multivariate indices and models of community and ecosystem function. However, the utility of trait-based approaches in ecology will benefit from efforts that demonstrate how these traits and indices influence organismal, community, and ecosystem processes across vegetation types, which may be achieved through meta-analysis and enhancement of trait databases. Additionally, intraspecific trait variation and species interactions need to be incorporated into predictive models using tools such as Bayesian hierarchical modelling. Finally, existing models linking traits to community and ecosystem processes need to be empirically tested for their applicability to be realized.

Seedling root responses to soil moisture and the identification of a belowground trait spectrum across three growth forms.

Root trait variation and plasticity could be key factors differentiating plant performance under drought. However, water manipulation and root measurements are rarely coupled empirically across growth forms to identify whether belowground strategies are generalizable across species. We measured seedling root traits across three moisture levels in 18 Mediterranean forbs, grasses, and woody species. Drought increased the root mass fraction (RMF) and decreased the relative proportion of thin roots (indicated by increased root diameters and decreased specific root length (SRL)), rates of root elongation and growth, plant nitrogen uptake, and plant growth. Although responses varied across species, plasticity was not associated with growth form. Woody species differed from forbs and grasses in many traits, but herbaceous groups were similar. Across water treatments, trait correlations suggested a single spectrum of belowground trade-offs related to resource acquisition and plant growth. While effects of SRL and RMF on plant growth shifted with drought, root elongation rate consistently represented this spectrum. We demonstrate that general patterns of root morphology and plasticity are identifiable across diverse species. Root trait measurements should enhance our understanding of belowground strategy and performance across growth forms, but it will be critical to incorporate plasticity and additional aspects of root function into these efforts.

Do key dimensions of seed and seedling functional trait variation capture variation in recruitment probability?

Seedling recruitment is a critical driver of population dynamics and community assembly, yet we know little about functional traits that define different recruitment strategies. For the first time, we examined whether trait relatedness across germination and seedling stages allows the identification of general recruitment strategies which share core functional attributes and also correspond to recruitment outcomes in applied settings. We measured six seed and eight seedling traits (lab- and field-collected, respectively) for 47 varieties of dryland grasses and used principal component analysis (PCA) and cluster analysis to identify major dimensions of trait variation and to isolate trait-based recruitment groups, respectively. PCA highlighted some links between seed and seedling traits, suggesting that relative growth rate and root elongation rate are simultaneously but independently associated with seed mass and initial root mass (first axis), and with leaf dry matter content, specific leaf area, coleoptile tissue density and germination rate (second axis). Third and fourth axes captured separate tradeoffs between hydrothermal time and base water potential for germination, and between specific root length and root mass ratio, respectively. Cluster analysis separated six recruitment types along dimensions of germination and growth rates, but classifications did not correspond to patterns of germination, emergence or recruitment in the field under either of two watering treatments. Thus, while we have begun to identify major threads of functional variation across seed and seedling stages, our understanding of how this variation influences demographic processes-particularly germination and emergence-remains a key gap in functional ecology.