Constraints on coastal dune invasion for a notorious plant invader.

Although most biological invasions are not successful, relatively few studies have examined otherwise notorious invaders in systems where they are not highly problematic. The annual grass Bromus tectorum is a dominant invader in western North America, but is usually confined to human-dominated and disturbed systems (e.g. roadsides and parking lots) in the East where it remains virtually unstudied. This study aims to address fundamental ecological questions regarding B. tectorum in a Cape Cod dune ecosystem. (i) What is the range of variation in population dynamics and the potential for population growth? (ii) Which factors influence its local abundance and distribution? We observed substantial variation in population dynamics over 3 years, with the number of adult B. tectorum individuals increasing substantially between the first 2 years (λ = 9.24) and then decreasing (λ = 0.43). Population growth in terms of total seeds was similarly variable, but to a lesser extent (λ = 2.32 followed by λ = 0.32). Experimental soil disturbance led to a more than 10-fold increase in mean seedling emergence, and high sensitivity to differences in emergence carried this effect through the life cycle. In contrast, barriers to seed dispersal had no effect on population dynamics, suggesting limited dispersal in this system. Across the landscape, the presence of B. tectorum was associated with areas of higher plant diversity as opposed to those with a strong dominant (e.g. the foredune, dominated by Ammophila breviligulata, or low heathlands, characterized by Hudsonia tomentosa and Arctostaphylos uva-ursi). Overall, we find that B. tectorum is capable of both substantial population growth and decline in a dune ecosystem, but is likely limited without disturbance and dispersal agents. Thus, management actions that restrict dune access (e.g. for nesting habitat) likely have the co-benefit of limiting the invasive potential of B. tectorum.

Impact of intra- versus inter-annual snow depth variation on water relations and photosynthesis for two Great Basin Desert shrubs.

Snowfall provides the majority of soil water in certain ecosystems of North America. We tested the hypothesis that snow depth variation affects soil water content, which in turn drives water potential (Ψ) and photosynthesis, over 10 years for two widespread shrubs of the western USA. Stem Ψ (Ψ stem) and photosynthetic gas exchange [stomatal conductance to water vapor (g s), and CO2 assimilation (A)] were measured in mid-June each year from 2004 to 2013 for Artemisia tridentata var. vaseyana (Asteraceae) and Purshia tridentata (Rosaceae). Snow fences were used to create increased or decreased snow depth plots. Snow depth on +snow plots was about twice that of ambient plots in most years, and 20 % lower on -snow plots, consistent with several down-scaled climate model projections. Maximal soil water content at 40- and 100-cm depths was correlated with February snow depth. For both species, multivariate ANOVA (MANOVA) showed that Ψ stem, g s, and A were significantly affected by intra-annual variation in snow depth. Within years, MANOVA showed that only A was significantly affected by spatial snow depth treatments for A. tridentata, and Ψ stem was significantly affected by snow depth for P. tridentata. Results show that stem water relations and photosynthetic gas exchange for these two cold desert shrub species in mid-June were more affected by inter-annual variation in snow depth by comparison to within-year spatial variation in snow depth. The results highlight the potential importance of changes in inter-annual variation in snowfall for future shrub photosynthesis in the western Great Basin Desert.

Effects of climate and snow depth on Bromus tectorum population dynamics at high elevation.

Invasive plants are thought to be especially capable of range shifts or expansion in response to climate change due to high dispersal and colonization abilities. Although highly invasive throughout the Intermountain West, the presence and impact of the grass Bromus tectorum has been limited at higher elevations in the eastern Sierra Nevada, potentially due to extreme wintertime conditions. However, climate models project an upward elevational shift of climate regimes in the Sierra Nevada that could favor B. tectorum expansion. This research specifically examined the effects of experimental snow depth manipulations and interannual climate variability over 5 years on B. tectorum populations at high elevation (2,175 m). Experimentally-increased snow depth had an effect on phenology and biomass, but no effect on individual fecundity. Instead an experimentally-increased snowpack inhibited population growth in 1 year by reducing seedling emergence and early survival. A similar negative effect of increased snow was observed 2 years later. However, a strong negative effect on B. tectorum was also associated with a naturally low-snow winter, when seedling emergence was reduced by 86%. Across 5 years, winters with greater snow cover and a slower accumulation of degree-days coincided with higher B. tectorum seedling density and population growth. Thus, we observed negative effects associated with both experimentally-increased and naturally-decreased snowpacks. It is likely that the effect of snow at high elevation is nonlinear and differs from lower elevations where wintertime germination can be favorable. Additionally, we observed a doubling of population size in 1 year, which is alarming at this elevation.

Positive effects of native shrubs on Bromus tectorum demography.

There is increasing recognition that overall interactions among plant species are often the net result of both positive and negative effects. However, the positive influence of other plants has rarely been examined using detailed demographic methods, which are useful for partitioning net effects at the population level into positive and/or negative effects on individual vital rates. This study examines the influence of microhabitats created by the native shrubs Artemisia tridentata and Purshia tridentata on the demography of the invasive annual grass Bromus tectorum in the Great Basin Desert, California, USA. Shrub understory environments differed significantly from intershrub space and were characterized by higher soil fertility and less extreme microclimates. There existed a strong spatial association between B. tectorum and the shrubs across four years, with more than double the density of B. tectorum in shrub microhabitats compared to intershrub space. Periodic matrix models were used to calculate population growth (lamda) and reproductive potential (RP, expected lifetime fecundity of seedlings) of B. tectorum in different microhabitats over two years. Modeled population growth was significantly increased in shrub microhabitats in the first of two years. This was primarily due to increased seedling establishment in Artemisia microhabitats, rather than effects during the growing season. In the following year, B. tectorum individuals in shrub microhabitats had a significantly greater reproductive potential than those in intershrub microhabitats, indicating shrub facilitation during the growing season. Loop analysis revealed an interacting effect of year and microhabitat on B. tectorum life history pathway elasticity values, demonstrating a fundamental influence of spatiotemporal factors on which life history pathways are important and/or possible. Life table response experiment (LTRE) analysis showed that increased survival and growth rates positively contributed to population growth in both years under Purshia, but only in the second year under Artemisia. This research provides evidence that the positive effects of native shrubs on B. tectorum can be strong enough to produce net positive effects at the population level, although positive effects were variable. In this study, a rigorous demographic approach was particularly useful in partitioning overall interactions into positive and negative components.