Andrew R. Kleinhesselink

Responses to invasion and invader removal differ between native and exotic plant groups in a coastal dune

The spread of exotic, invasive species is a global phenomenon that is recognized as a major source of environmental change. Although many studies have addressed the effects of exotic plants on the communities they invade, few have quantified the effects of invader removal on plant communities, or considered the degree to which different plant groups vary in response to invasion and invader removal. We evaluated the effects of an exotic succulent, iceplant (Carpobrotus edulis), on a coastal dune plant community in northern California, as well as the community responses to its removal. To assess possible mechanisms by which iceplant affects other plants, we also evaluated its above- and belowground influences on the germination and growth of a dominant exotic annual grass, Bromus diandrus. We found that iceplant invasion was associated with reduced native plant cover as well as increased cover and density of some exotic plants—especially exotic annual grasses. However, iceplant removal did not necessarily lead to a reversal of these effects: removal increased the cover and density of both native and exotic species. We also found that B. diandrus grown in iceplant patches, or in soil where iceplant had been removed, had poorer germination and growth than B. diandrus grown in soil not influenced by iceplant. This suggests that the influence of iceplant on this dune plant community occurs, at least in part, due to belowground effects, and that these effects remain after iceplant has been removed. Our study demonstrates the importance of considering how exotic invasive plants affect not only native species, but also co-occurring exotic taxa. It also shows that combining observational studies with removal experiments can lead to important insights into the influence of invaders and the mechanisms of their effects.

Indirect Effects of Environmental Change in Resource Competition Models.

Anthropogenic environmental change can affect species directly by altering physiological rates or indirectly by changing competitive outcomes. The unknown strength of competition-mediated indirect effects makes it difficult to predict species abundances in the face of ongoing environmental change. Theory developed with phenomenological competition models shows that indirect effects are weak when coexistence is strongly stabilized, but these models lack a mechanistic link between environmental change and species performance. To extend existing theory, we examined the relationship between coexistence and indirect effects in mechanistic resource competition models. We defined environmental change as a change in resource supply points and quantified the resulting competition-mediated indirect effects on species abundances. We found that the magnitude of indirect effects increases in proportion to niche overlap. However, indirect effects also depend on differences in how competitors respond to the change in resource supply, an insight hidden in nonmechanistic models. Our analysis demonstrates the value of using niche overlap to predict the strength of indirect effects and clarifies the types of indirect effects that global change can have on competing species.

Direct effects dominate responses to climate perturbations in grassland plant communities

Theory predicts that strong indirect effects of environmental change will impact communities when niche differences between competitors are small and variation in the direct effects experienced by competitors is large, but empirical tests are lacking. Here we estimate negative frequency dependence, a proxy for niche differences, and quantify the direct and indirect effects of climate change on each species. Consistent with theory, in four of five communities indirect effects are strongest for species showing weak negative frequency dependence. Indirect effects are also stronger in communities where there is greater variation in direct effects. Overall responses to climate perturbations are driven primarily by direct effects, suggesting that single species models may be adequate for forecasting the impacts of climate change in these communities.

Shrubs as ecosystem engineers across an environmental gradient: effects on species richness and exotic plant invasion

Ecosystem-engineering plants modify the physical environment and can increase species diversity and exotic species invasion. At the individual level, the effects of ecosystem engineers on other plants often become more positive in stressful environments. In this study, we investigated whether the community-level effects of ecosystem engineers also become stronger in more stressful environments. Using comparative and experimental approaches, we assessed the ability of a native shrub (Ericameria ericoides) to act as an ecosystem engineer across a stress gradient in a coastal dune in northern California, USA. We found increased coarse organic matter and lower wind speeds within shrub patches. Growth of a dominant invasive grass (Bromus diandrus) was facilitated both by aboveground shrub biomass and by growing in soil taken from shrub patches. Experimental removal of shrubs negatively affected species most associated with shrubs and positively affected species most often found outside of shrubs. Counter to the stress-gradient hypothesis, the effects of shrubs on the physical environment and individual plant growth did not increase across the established stress gradient at this site. At the community level, shrub patches increased beta diversity, and contained greater rarified richness and exotic plant cover than shrub-free patches. Shrub effects on rarified richness increased with environmental stress, but effects on exotic cover and beta diversity did not. Our study provides evidence for the community-level effects of shrubs as ecosystem engineers in this system, but shows that these effects do not necessarily become stronger in more stressful environments.

Ecosystem functional response across precipitation extremes in a sagebrush steppe

Background Precipitation is predicted to become more variable in the western United States, meaning years of above and below average precipitation will become more common. Periods of extreme precipitation are major drivers of interannual variability in ecosystem functioning in water limited communities, but how ecosystems respond to these extremes over the long-term may shift with precipitation means and variances. Long-term changes in ecosystem functional response could reflect compensatory changes in species composition or species reaching physiological thresholds at extreme precipitation levels. Methods We conducted a five year precipitation manipulation experiment in a sagebrush steppe ecosystem in Idaho, United States. We used drought and irrigation treatments (approximately 50% decrease/increase) to investigate whether ecosystem functional response remains consistent under sustained high or low precipitation. We recorded data on aboveground net primary productivity (ANPP), species abundance, and soil moisture. We fit a generalized linear mixed effects model to determine if the relationship between ANPP and soil moisture differed among treatments. We used nonmetric multidimensional scaling to quantify community composition over the five years. Results Ecosystem functional response, defined as the relationship between soil moisture and ANPP, was similar among irrigation and control treatments, but the drought treatment had a greater slope than the control treatment. However, all estimates for the effect of soil moisture on ANPP overlapped zero, indicating the relationship is weak and uncertain regardless of treatment. There was also large spatial variation in ANPP within-years, which contributes to the uncertainty of the soil moisture effect. Plant community composition was remarkably stable over the course of the experiment and did not differ among treatments. Discussion Despite some evidence that ecosystem functional response became more sensitive under sustained drought conditions, the response of ANPP to soil moisture was consistently weak and community composition was stable. The similarity of ecosystem functional responses across treatments was not related to compensatory shifts at the plant community level, but instead may reflect the insensitivity of the dominant species to soil moisture. These species may be successful precisely because they have evolved life history strategies that buffer them against precipitation variability.