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Understanding the resistance and resilience of foundation plant species to climate change is a critical issue because the loss of these species would fundamentally reshape communities and ecosystem processes. High levels of population genetic diversity may buffer foundation species against climate disruptions, but the strong selective pressures associated with climatic shifts may also rapidly reduce such diversity. We characterized genetic diversity and its responsiveness to experimental drought in the foundation plant, black grama grass (Bouteloua eriopoda), which dominates many western North American grasslands and shrublands. Previous studies suggested that in arid ecosystems, black grama reproduces largely asexually via stolons, and thus is likely to have low genetic variability, which might limit its potential to respond to climate disruptions. Using genotyping-by-sequencing, we demonstrated unexpectedly high genetic variability among black grama plants in a 1 ha site within the Sevilleta National Wildlife Refuge in central New Mexico, suggesting some level of sexual reproduction. Three years of experimental, growing season drought reduced black grama survival and biomass (the latter by 96%), with clear genetic differentiation (higher F ST ) between plants succumbing to drought and those remaining alive. Reduced genetic variability in the surviving plants in drought plots indicated that the experimental drought had forced black grama populations through selection bottlenecks. These results suggest that foundation grass species, such as black grama, may experience rapid evolutionary change if future climates include more severe droughts.

For the southwestern United States, climate models project an increase in extreme precipitation events and prolonged dry periods. While most studies emphasize plant functional type response to precipitation variability, it is also important to understand the physiological characteristics of dominant plant species that define plant community composition and, in part, regulate ecosystem response to climate change. We utilized rainout shelters to alter the magnitude and frequency of rainfall and measured the physiological response of the dominant C4 grasses, Bouteloua eriopoda and Bouteloua gracilis. We hypothesized that: (1) the more drought-adapted B. eriopoda would exhibit faster recovery and higher rates of leaf-level photosynthesis (A(net)) than B. gracilis, (2) A(net) would be greater under the higher average soil water content in plots receiving 30-mm rainfall events, (3) co-dominance of B. eriopoda and B. gracilis in the ecotone would lead to intra-specific differences from the performance of each species at the site where it was dominant. Throughout the study, soil moisture explained 40-70% of the variation in A(net). Consequently, differences in rainfall treatments were not evident from intra-specific physiological function without sufficient divergence in soil moisture. Under low frequency, larger rainfall events B. gracilis exhibited improved water status and longer periods of C gain than B. eriopoda. Results from this study indicate that less frequent and larger rainfall events could provide a competitive advantage to B. gracilis and influence species composition across this arid-semiarid grassland ecotone.

Aridland ecosystems are predicted to be responsive to both increases and decreases in precipitation. In addition, chronic droughts may contribute to encroachment of native C3 shrubs into C4-dominated grasslands. We conducted a long-term rainfall manipulation experiment in native grassland, shrubland and the grass-shrub ecotone in the northern Chihuahuan Desert, USA. We evaluated the effects of 5 years of experimental drought and 4 years of water addition on plant community structure and dynamics. We assessed the effects of altered rainfall regimes on the abundance of dominant species as well as on species richness and subdominant grasses, forbs and shrubs. Nonmetric multidimensional scaling and MANOVA were used to quantify changes in species composition in response to chronic addition or reduction of rainfall. We found that drought consistently and strongly decreased cover of Bouteloua eriopoda, the dominant C4 grass in this system, whereas water addition slightly increased cover, with little variation between years. In contrast, neither chronic drought nor increased rainfall had consistent effects on the cover of Larrea tridentata, the dominant C3 shrub. Species richness declined in shrub-dominated vegetation in response to drought whereas richness increased or was unaffected by water addition or drought in mixed- and grass-dominated vegetation. Cover of subdominant shrubs, grasses and forbs changed significantly over time, primarily in response to interannual rainfall variability more so than to our experimental rainfall treatments. Nevertheless, drought and water addition shifted the species composition of plant communities in all three vegetation types. Overall, we found that B. eriopoda responded strongly to drought and less so to irrigation, whereas L. tridentata showed limited response to either treatment. The strong decline in grass cover and the resistance of shrub cover to rainfall reduction suggest that chronic drought may be a key factor promoting shrub dominance during encroachment into desert grassland.

Variation in freezing severity is hypothesized to have influenced the distribution and evolution of the warm desert evergreen genus Larrea. If this hypothesis is correct, performance and survival of species and populations should vary predictably along gradients of freezing severity. If freezing environment changes in the future, the ability of Larrea to adapt will depend on the structure of variation for freezing resistance within populations. To test whether freezing responses vary among and within Larrea populations, we grew maternal families of seedlings from high and low latitude L. divaricata and high latitude L. tridentata populations in a common garden. We measured survival, projected plant area and dark-adapted chlorophyll fluorescence (F (v) /F (m)) before and after cold acclimation and for 2 weeks following a single freeze. We detected significant variation in freezing resistance among species and populations. Maternal family lines differed significantly in their responses to cold acclimation and/or freezing for two out of the three populations: among L. tridentata maternal families and among low latitude L. divaricata maternal families. There were no significant differences across maternal families of high latitude L. divaricata. Our results indicate that increased freezing resistance in high latitude populations likely facilitated historical population expansion of both species into colder climates, but this may have occurred to a greater extent for L. tridentata than for L. divaricata. Differences in the structure of variation for cold acclimation and freezing responses among populations suggest potential differences in their ability to evolve in response to future changes in freezing severity.

In the arid and semiarid regions of North America, discrete precipitation pulses are important triggers for biological activity. The timing and magnitude of these pulses may differentially affect the activity of plants and microbes, combining to influence the C balance of desert ecosystems. Here, we evaluate how a "pulse" of water influences physiological activity in plants, soils and ecosystems, and how characteristics, such as precipitation pulse size and frequency are important controllers of biological and physical processes in arid land ecosystems. We show that pulse size regulates C balance by determining the temporal duration of activity for different components of the biota. Microbial respiration responds to very small events, but the relationship between pulse size and duration of activity likely saturates at moderate event sizes. Photosynthetic activity of vascular plants generally increases following relatively larger pulses or a series of small pulses. In this case, the duration of physiological activity is an increasing function of pulse size up to events that are infrequent in these hydroclimatological regions. This differential responsiveness of photosynthesis and respiration results in arid ecosystems acting as immediate C sources to the atmosphere following rainfall, with subsequent periods of C accumulation should pulse size be sufficient to initiate vascular plant activity. Using the average pulse size distributions in the North American deserts, a simple modeling exercise shows that net ecosystem exchange of CO2 is sensitive to changes in the event size distribution representative of wet and dry years. An important regulator of the pulse response is initial soil and canopy conditions and the physical structuring of bare soil and beneath canopy patches on the landscape. Initial condition influences responses to pulses of varying magnitude, while bare soil/beneath canopy patches interact to introduce nonlinearity in the relationship between pulse size and soil water response. Building on this conceptual framework and developing a greater understanding of the complexities of these eco-hydrologic systems may enhance our ability to describe the ecology of desert ecosystems and their sensitivity to global change.

Wood density (D t ), an excellent predictor of mechanical properties, is typically viewed in relation to support against gravity, wind, snow, and other environmental forces. In contrast, we show the surprising extent to which variation in D t and wood structure is linked to support against implosion by negative pressure in the xylem pipeline. The more drought-tolerant the plant, the more negative the xylem pressure can become without cavitation, and the greater the internal load on the xylem conduit walls. Accordingly, D t was correlated with cavitation resistance. This trend was consistent with the maintenance of a safety factor from implosion by negative pressure: conduit wall span (b) and thickness (t) scaled so that (t/b) 2 was proportional to cavitation resistance as required to avoid wall collapse. Unexpectedly, trends in D t may be as much or more related to support of the xylem pipeline as to support of the plant.