Seedling responses to water pulses in shrubs with contrasting histories of grassland encroachment.

Woody plant encroachment into grasslands has occurred worldwide, but it is unclear why some tree and shrub species have been markedly more successful than others. For example, Prosopis velutina has proliferated in many grasslands of the Sonoran Desert in North America over the past century, while other shrub species with similar growth form and life history, such as Acacia greggii, have not. We conducted a glasshouse experiment to assess whether differences in early seedling development could help explain why one species and not the other came to dominate many Sonoran Desert grasslands. We established eight watering treatments mimicking a range of natural precipitation patterns and harvested seedlings 16 or 17 days after germination. A. greggii had nearly 7 times more seed mass than P. velutina, but P. velutina emerged earlier (by 3.0±0.3 d) and grew faster (by 8.7±0.5 mg d⁻¹). Shoot mass at harvest was higher in A. greggii (99±6 mg seedling⁻¹) than in P. velutina (74±2 mg seedling⁻¹), but there was no significant difference in root mass (54±3 and 49±2 mg seedling⁻¹, respectively). Taproot elongation was differentially sensitive to water supply: under the highest initial watering pulse, taproots were 52±19 mm longer in P. velutina than in A. greggii. Enhanced taproot elongation under favorable rainfall conditions could give nascent P. velutina seedlings growth and survivorship advantages by helping reduce competition with grasses and maintain contact with soil water during drought. Conversely, A. greggii's greater investment in mass per seed appeared to provide little return in early seedling growth. We suggest that such differences in recruitment traits and their sensitivities to environmental conditions may help explain ecological differences between species that are highly similar as adults and help identify pivotal drivers of shrub encroachment into grasslands.

Precipitation pulses and carbon fluxes in semiarid and arid ecosystems.

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.

Convergence across biomes to a common rain-use efficiency.

Water availability limits plant growth and production in almost all terrestrial ecosystems. However, biomes differ substantially in sensitivity of aboveground net primary production (ANPP) to between-year variation in precipitation. Average rain-use efficiency (RUE; ANPP/precipitation) also varies between biomes, supposedly because of differences in vegetation structure and/or biogeochemical constraints. Here we show that RUE decreases across biomes as mean annual precipitation increases. However, during the driest years at each site, there is convergence to a common maximum RUE (RUE(max)) that is typical of arid ecosystems. RUE(max) was also identified by experimentally altering the degree of limitation by water and other resources. Thus, in years when water is most limiting, deserts, grasslands and forests all exhibit the same rate of biomass production per unit rainfall, despite differences in physiognomy and site-level RUE. Global climate models predict increased between-year variability in precipitation, more frequent extreme drought events, and changes in temperature. Forecasts of future ecosystem behaviour should take into account this convergent feature of terrestrial biomes.

Resource pulses, species interactions, and diversity maintenance in arid and semi-arid environments.

Arid environments are characterized by limited and variable rainfall that supplies resources in pulses. Resource pulsing is a special form of environmental variation, and the general theory of coexistence in variable environments suggests specific mechanisms by which rainfall variability might contribute to the maintenance of high species diversity in arid ecosystems. In this review, we discuss physiological, morphological, and life-history traits that facilitate plant survival and growth in strongly water-limited variable environments, outlining how species differences in these traits may promote diversity. Our analysis emphasizes that the variability of pulsed environments does not reduce the importance of species interactions in structuring communities, but instead provides axes of ecological differentiation between species that facilitate their coexistence. Pulses of rainfall also influence higher trophic levels and entire food webs. Better understanding of how rainfall affects the diversity, species composition, and dynamics of arid environments can contribute to solving environmental problems stemming from land use and global climate change.