Sarah C. Elmendorf

RAPID EVOLUTION OF AN INVASIVE PLANT

Exotic plants often face different conditions from those experienced where they are native. The general issue of how exotics respond to unfamiliar environments within their new range is not well understood. Phenotypic plasticity has historically been seen as the primary mechanism enabling exotics to colonize large, environmentally diverse areas. However, new work indicates that exotics can evolve quickly, suggesting that contemporary evolution may be more important in invasion ecology than previously appreciated. To determine the influence of contemporary evolution, phenotypic plasticity, and founder ef- fects in affecting phenotypic variation among introduced plants, we compared the size, fecundity, and leaf area of St. Johns wort ( Hypericum perforatum) collected from native European and introduced western and central North American populations in common gardens in Washington, California, Spain, and Sweden. We also determined genetic rela- tionships among these plants by examining variation in amplified fragment length poly- morphism (AFLP) markers. There was substantial genetic variation among introduced populations and evidence for multiple introductions of H. perforatum into North America. Across common gardens in- troduced plants were neither universally larger nor more fecund than natives. However, within common gardens, both introduced and native populations exhibited significant lat- itudinally based clines in size and fecundity. Clines among introduced populations broadly converged with those among native populations. Introduced and native plants originating from northern latitudes generally outperformed those originating from southern latitudes when grown in northern latitude gardens of Washington and Sweden. Conversely, plants from southern latitudes performed best in southern gardens in Spain and California. Clinal patterns in leaf area, however, did not change between gardens; European and central North American plants from northern latitudes had larger leaves than plants from southern latitudes within these regions in both Washington and California, the two gardens where this trait was measured. Introduced plants did not always occur at similar latitudes as their most closely related native progenitor, indicating that pre-adaptation (i.e., climate matching) is unlikely to be the sole explanation for clinal patterns among introduced populations. Instead, results suggest that introduced plants are evolving adaptations to broad-scale environmental conditions in their introduced range.

AN INTRODUCED PREDATOR ALTERS ALEUTIAN ISLAND PLANT COMMUNITIES BY THWARTING NUTRIENT SUBSIDIES

The ramifying effects of top predators on food webs traditionally have been studied within the framework of trophic cascades. Trophic cascades are compelling because they embody powerful indirect effects of predators on primary production. Although less studied, indirect effects of predators may occur via routes that are not exclusively trophic. We quantified how the introduction of foxes onto the Aleutian Islands transformed plant communities by reducing abundant seabird populations, thereby disrupting nutrient sub- sidies vectored by seabirds from sea to land. We compared soil and plant fertility, plant biomass and community composition, and stable isotopes of nitrogen in soil, plants, and other organisms on nine fox-infested and nine historically fox-free islands across the Aleu- tians. Additionally, we experimentally augmented nutrients on a fox-infested island to test whether differences in plant productivity and composition between fox-infested and fox- free islands could have arisen from differences in nutrient inputs between island types. Islands with historical fox infestations had soils low in phosphorus and nitrogen and plants low in tissue nitrogen. Soils, plants, slugs, flies, spiders, and bird droppings on these islands had low d 15 N values indicating that these organisms obtained nitrogen from internally derived sources. In contrast, soils, plants, and higher trophic level organisms on fox-free islands had elevated d 15 N signatures indicating that they utilized nutrients derived from the marine environment. Furthermore, soil phosphorus (but not nitrogen) and plant tissue ni- trogen were higher on fox-free than fox-infested islands. Nutrient subsidized fox-free islands supported lush, high biomass plant communities dominated by graminoids. Fox-infested islands were less graminoid dominated and had higher cover and biomass of low-lying forbs and dwarf shrubs. While d 15 N profiles of soils and plants and graminoid biomass varied with island size and distance from shore, after accounting for these effects differences between fox-infested and fox-free islands still existed. Fertilization over four years caused a 24-fold increase in graminoid biomass and a shift toward a more graminoid dominated plant community typical of fox-free islands. These results indicate that apex predators can influence plant productivity and composition through complex interaction web pathways involving both top-down forcing and bottom-up nutrient exchanges across systems.

CONTRASTING PLANT PHYSIOLOGICAL ADAPTATION TO CLIMATE IN THE NATIVE AND INTRODUCED RANGE OF HYPERICUM PERFORATUM

Abstract How introduced plants, which may be locally adapted to specific climatic conditions in their native range, cope with the new abiotic conditions that they encounter as exotics is not well understood. In particular, it is unclear what role plasticity versus adaptive evolution plays in enabling exotics to persist under new environmental circumstances in the introduced range. We determined the extent to which native and introduced populations of St. Johns Wort (Hypericum perforatum) are genetically differentiated with respect to leaf-level morphological and physiological traits that allow plants to tolerate different climatic conditions. In common gardens in Washington and Spain, and in a greenhouse, we examined clinal variation in percent leaf nitrogen and carbon, leaf δ13C values (as an integrative measure of water use efficiency), specific leaf area (SLA), root and shoot biomass, root/shoot ratio, total leaf area, and leaf area ratio (LAR). As well, we determined whether native European H. perforatum experienced directional selection on leaf-level traits in the introduced range and we compared, across gardens, levels of plasticity in these traits. In field gardens in both Washington and Spain, native populations formed latitudinal clines in percent leaf N. In the greenhouse, native populations formed latitudinal clines in root and shoot biomass and total leaf area, and in the Washington garden only, native populations also exhibited latitudinal clines in percent leaf C and leaf δ13C. Traits that failed to show consistent latitudinal clines instead exhibited significant phenotypic plasticity. Introduced St. Johns Wort populations also formed significant or marginally significant latitudinal clines in percent leaf N in Washington and Spain, percent leaf C in Washington, and in root biomass and total leaf area in the greenhouse. In the Washington common garden, there was strong directional selection among European populations for higher percent leaf N and leaf δ13C, but no selection on any other measured trait. The presence of convergent, genetically based latitudinal clines between native and introduced H. perforatum, together with previously published molecular data, suggest that native and exotic genotypes have independently adapted to a broad-scale variation in climate that varies with latitude.

Temporal variability and nestedness in California grassland species composition.

Nestedness occurs when species-poor assemblages contain a subset of the species that occur in more species-rich communities and is a commonly observed pattern in spatial data sets. Examination of nested distribution patterns across time rather than space are rarely conducted, even though they may have important implications for species coexistence. Nested temporal assemblages can occur when most species respond similarly to interannual variation in conditions. In contrast, assemblages might be non-nested when different sets of species occur in different years, either because of different resource requirements or as a result of competitive exclusion. Using eight years of plant occurrence data at 71 sites in California grasslands, we found strong signals of temporal nestedness with most species favored by similar conditions. High-quality years enabled the expansion of both grasses and forbs into locales where they were not found during poor-quality years. Native annual forb, exotic annual forb, and exotic annual grass species richness were all greatest in cool, wet years following hot, dry years. Together, these analyses support the hypothesis that, in the absence of community members that specialize on poor-quality years, interannual environmental variation can cause communities to form nested subsets across time much as they do across space.

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.