Benchmarking novel approaches for modelling species range dynamics.

Increasing biodiversity loss due to climate change is one of the most vital challenges of the 21st century. To anticipate and mitigate biodiversity loss, models are needed that reliably project species' range dynamics and extinction risks. Recently, several new approaches to model range dynamics have been developed to supplement correlative species distribution models (SDMs), but applications clearly lag behind model development. Indeed, no comparative analysis has been performed to evaluate their performance. Here, we build on process-based, simulated data for benchmarking five range (dynamic) models of varying complexity including classical SDMs, SDMs coupled with simple dispersal or more complex population dynamic models (SDM hybrids), and a hierarchical Bayesian process-based dynamic range model (DRM). We specifically test the effects of demographic and community processes on model predictive performance. Under current climate, DRMs performed best, although only marginally. Under climate change, predictive performance varied considerably, with no clear winners. Yet, all range dynamic models improved predictions under climate change substantially compared to purely correlative SDMs, and the population dynamic models also predicted reasonable extinction risks for most scenarios. When benchmarking data were simulated with more complex demographic and community processes, simple SDM hybrids including only dispersal often proved most reliable. Finally, we found that structural decisions during model building can have great impact on model accuracy, but prior system knowledge on important processes can reduce these uncertainties considerably. Our results reassure the clear merit in using dynamic approaches for modelling species' response to climate change but also emphasize several needs for further model and data improvement. We propose and discuss perspectives for improving range projections through combination of multiple models and for making these approaches operational for large numbers of species.

Can spatial isolation help predict dispersal-limited sites for native species restoration?

When the distribution of species is limited by propagule supply, new populations may be initiated by seed addition, but identifying suitable sites for efficiently targeted seed addition remains a major challenge for restoration. In addition to the biotic or abiotic variables typically used in species distribution models, spatial isolation from conspecifics could help predict the suitability of unoccupied sites. Site suitability might be expected to increase with spatial isolation after other factors are accounted for, since isolation increases the chance that a site is unoccupied only because of propagule limitation. For two native annual forbs in Californian grasslands, we combined experimental seeding and niche modeling to ask whether suitability of unoccupied sites could be predicted by spatial variables (either distances from, or densities of, conspecific populations), either by themselves or in combination with niche models. We also asked whether experimental tests of these predictions held up not only in the short term (one year), but also in the longer term (three years). For Lasthenia californica, seed additions were only successful relatively near existing populations. For Lupinus nanus, seeding success was low and was positively related to the number of conspecifics within 1 km. For both species, a few previously unoccupied sites remained occupied three years after seeding, but this subset was not predictable based on either spatial or niche variables. Seed addition alone may be a limited means of native forb restoration if suitable unoccupied sites are either rare or unpredictable, or if they tend to be close to where the species already occurs.

Spatial niches and coexistence: testing theory with tarweeds.

Competitive coexistence in a spatially heterogeneous environment is traditionally attributed to niche differences, but several recent theories have proposed important additional roles for propagule limitation and chance (e.g., neutral theory, stochastic niche theory, spatial storage effect). We tested whether propagule supply and timing of disturbance affected the coexistence of three ecologically similar plants that replace one another with partial overlap along a local soil gradient. We asked what prevents the species that dominates the most common habitat (Holocarpha virgata, open hillsides) from invading the habitats where the other two species are dominant (Calycadenia pauciflora, rocky hilltops; Hemizonia congesta, clay-rich bottomlands). We added abundant Holocarpha seeds into Calycadenia and Hemizonia habitats that were experimentally disturbed at different times of year. Initial Holocarpha seedling densities in Calycadenia and Hemizonia habitats equaled or exceeded those in unmanipulated Holocarpha habitat, but Holocarpha survival, adult size, and fecundity were much lower outside its own habitat. Holocarpha persisted in Calycadenia and Hemizonia habitats for three years, and springtime disturbance promoted this expansion. However, outside its own habitat Holocarpha showed below-replacement fitness and little competitive effect on the other two species. Our results were most consistent with a deterministic view of spatial niches. Nonetheless, chance events may often cause natural communities to include some transient populations at any given time, leading them to appear "unsaturated" with species.

Fluctuating patch boundaries in a native annual forb: the roles of niche and dispersal limitation.

Abiotic, biotic, and dispersal constraints jointly control the spatial distributions of species, but few studies have directly evaluated how these forces interact and vary over time to create dynamic spatial distributions. Through three years of observation and two years of field manipulation, I investigated simultaneous constraints on the spatial distribution of Lupinus nanus, a native annual legume that grows in dense patches in California grasslands. I transplanted L. nanus across its own patch boundaries, yet within apparently suitable habitat, and assessed the demographic success of naturally occurring and seeded plants in plots with and without competitor removal in years that varied in temperature and rainfall. Core sites were defined as those consistently densely occupied, whereas peripheral sites were densely occupied only during some years, and exterior sites were consistently unoccupied or very sparsely occupied. Site types (core, periphery, and exterior) differed in soil moisture, P, and NO3. Competition limited emergence in all site types in the dry/warm year and in patch peripheries in the wet/cool year. Population fitness (seeds produced per seed added) was > 1.0 in cores during all years. Peripheral sites had fitness near replacement in the wet/cool year, which was greatly increased by competition removal. Exterior fitness was < 1.0 in both experimental years, regardless of seed addition and competitor removal. Seed addition did not increase site-specific fitness, and a seed bank was found to be present in all site types. Herbivory was greater in patch cores and peripheries than in exteriors. Soil variation exerted the most consistent control over patch limits, while competition played an intermittent role in excluding Lupinus from patch peripheries. The dynamic distribution of L. nanus is the product of temporal variation in specific abiotic and biotic niche axes, primarily soil characteristics and competition, rather than dispersal limitation.

Use of community-composition data to predict the fecundity and abundance of species.

Species distribution models are critical tools for the prediction of invasive species spread and conservation of biodiversity. The majority of species distribution models have been built with environmental data. Community ecology theory suggests that species co-occurrence data could also be used to predict current and potential distributions of species. Species assemblages are the products of biotic and environmental constraints on the distribution of individual species and as a result may contain valuable information for niche modeling. We compared the predictive ability of distribution models of annual grassland plants derived from either environmental or community-composition data. Composition-based models were built with the presence or absence of species at a site as predictors of site quality, whereas environment-based models were built with soil chemistry, moisture content, above-ground biomass, and solar radiation as predictors. The reproductive output of experimentally seeded individuals of 4 species and the abundance of 100 species were used to evaluate the resulting models. Community-composition data were the best predictors of both the site-specific reproductive output of sown individuals and the site-specific abundance of existing populations. Successful community-based models were robust to omission of data on the occurrence of rare species, which suggests that even very basic survey data on the occurrence of common species may be adequate for generating such models. Our results highlight the need for increased public availability of ecological survey data to facilitate community-based modeling at scales relevant to conservation.

Plant competition varies with community composition in an edaphically complex landscape.

There is currently no consensus on how physical and biological factors affect competitive intensity. Tests of whether competitive intensity varies along axes of environmental change have commonly been conducted in systems with a single strong environmental gradient, such as productivity, a soil resource, or an environmental stress. Frequently, these same axes are associated with changes in species composition, yet few studies have asked whether shifts in the identity of competitors affect competitive intensity. We ask whether resources (nutrients, water), stressors (heavy metals, Ca:Mg ratio), productivity (aboveground biomass), or species identity (an ordination axis of plant community composition) were the best predictors of the intensity of competition in a heterogeneous grassland landscape that included multiple independent environmental gradients. The reproductive fitness of six annual plant species was measured in the presence and absence of competitors and used to calculate relative interaction intensity (RII). We found that RII was best predicted by community composition. Nutrient availability was also important, and a post hoc test showed that competitive intensity was best explained by the combined effects of community composition and nutrient availability. We argue that community composition may be the most effective metric for predicting competitive intensity in many ecosystems because it includes both the competitive effects of the local community and information about covarying environmental characteristics.

Propagule vs. niche limitation: untangling the mechanisms behind plant species' distributions.

Distinguishing the roles of propagule limitation and niche requirements in controlling plant species distributions is important for understanding community structure, invasion, and restoration. We used species distribution models based on plant and environmental survey data to assess the strength of species' affinities for particular environmental conditions. We hypothesized that species with statistically detectable environmental requirements were primarily niche-limited, while species with weak habitat affinities were primarily propagule-limited. We tested this hypothesis via a seeding experiment in which we compared species' reproductive fitness in occupied and unoccupied sites. Species that appeared to be niche-limited based on distribution models had lower fitness when planted in unoccupied sites, while species that models suggested were propagule-limited had equivalent fitness when planted in occupied and unoccupied sites. Our results demonstrate that within a single community, both species limited primarily by niche availability or primarily by propagule availability can be identified using observational data.

Propagule limitation, disparate habitat quality, and variation in phenotypic selection at a local species range boundary.

Adaptation to novel conditions beyond current range boundaries requires the presence of suitable sites within dispersal range, but may be impeded when emigrants encounter poor habitat and sharply different selection pressures. We investigated fine-scale spatial heterogeneity in ecological dynamics and selection at a local population boundary of the annual plant Gilia tricolor. In two years, we planted G. tricolor seeds in core habitat, margin habitat at the edge of the local range, and exterior habitat in order to measure spatial and temporal variation in habitat quality, opportunity for selection, and selection on phenotypic traits. We found a striking decline in average habitat quality with distance from the population core, yet some migrant seeds were successful in suitable, unoccupied microsites at and beyond the range boundary. Total and direct selection on four out of five measured phenotypic traits varied across habitat zones, as well as between years. Moreover, the margin habitat often exerted unique selection pressures that were not intermediate between core and exterior habitats. This study reveals that a combination of ecological and evolutionary forces, including propagule limitation, variation in habitat quality and spatial heterogeneity in phenotypic selection may reduce opportunities for adaptive range expansion, even across a very local population boundary.