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.

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Seasonal changes in the hydrogen isotope ratios of xylem waters were measured to determine water sources used for growth in desert plants of southern Utah. While all species used winter-spring recharge precipitation for spring growth, utilization of summer rains was life-form dependent. Annuals and succulent perennials exhibited a complete dependence on summer precipitation. Herbaceous and woody perennial species simultaneously utilized both summer precipitation and remaining winter-spring precipitation, with herbaceous species much more reliant on the summer precipitation component. Several of the woody perennials exhibited no response to summer precipitation. Currently, precipitation in southern Utah is evenly partitioned between winter and summer time periods; however, global circulation models predict that summer precipitation will increase in response to anticipated climate change. Our data indicate that components within the community will differentially responde to the change in precipitation patterns. These results are discussed in relation to impact on competition and possible changes in community structure.

δ13C values for freshwater aquatic plant matter varies from −11 to −50‰ and is not a clear indicator of photosynthetic pathway as in terrestrial plants. Several factors affect δ13C of aquatic plant matter. These include: (1) The δ13C signature of the source carbon has been observed to range from +1‰ for HCO3− derived from limestone to −30‰ for CO2 derived from respiration. (2) Some plants assimilate HCO3−, which is –7 to –11‰ less negative than CO2. (3) C3, C4, and CAM photosynthetic pathways are present in aquatic plants. (4) Diffusional resistances are orders of magnitude greater in the aquatic environment than in the aerial environment. The greater viscosity of water acts to reduce mixing of the carbon pool in the boundary layer with that of the bulk solution. In effect, many aquatic plants draw from a finite carbon pool, and as in terrestrial plants growing in a closed system, biochemical discrimination is reduced. In standing water, this factor results in most aquatic plants having a δ13C value similar to the source carbon. Using Farquhar's equation and other physiological data, it is possible to use δ13C values to evaluate various parameters affecting photosynthesis, such as limitations imposed by CO2 diffusion and carbon source.

A number of operationally defined methods exist for pretreating plant tissues in order to measure C, N, and O isotopes. Because these isotope measurements are used to infer information about environmental conditions that existed at the time of tissue growth, it is important that these pretreatments remove compounds that may have exchanged isotopes or have been synthesized after the original formation of these tissues. In stable isotope studies, many pretreatment methods focus on isolating "cellulose" from the bulk tissue sample because cellulose does not exchange C and O isotopes after original synthesis. We investigated the efficacy of three commonly applied pretreatment methods, the Brendel method and two variants of the Brendel method, the Jayme-Wise method and successive acid/base/acid washes, for use on three tissue types (wood, leaves, roots). We then compared the effect of each method on C and O isotope composition ( 13 C, 14 C, 18 O), C and N content, and chemical composition of the residue produced (using 13 C nuclear magnetic resonance (NMR)). Our results raised concerns over use of the Brendel method as published, as it both added C and N to the sample and left a residue that contains remnant lipids and waxes. Furthermore, this method resulted in 18 O values that are enriched relative to the other methods. Modifying the Brendel method by adding a NaOH step (wash) solved many of these problems. We also found that processed residues vary by tissue type. For wood and root tissues, the 13 C NMR spectra and the 18 O and 13 C data showed only small differences between residues for the Jayme-Wise and

Although the destruction of tropical rain forests receives much attention, tropical dry forests are in general far more threatened and endangered. Eliminating grazing ungulates is often considered a key first step toward protecting these ecosystems, but few studies have investigated the long‐term effects of this technique. We examined the effects of ungulate exclusion from a 2.3‐ha native dry‐forest preserve on the island of Hawaii by comparing its present flora to the flora of an adjacent area subjected to continuous grazing since the preserve was fenced over 40 years ago. Relative to this adjacent area, the fenced preserve contained a more diverse flora with substantially greater coverage of native overstory and understory species. Until recently, however, regeneration of native canopy trees within the preserve appears to have been thwarted by a dominant herbaceous cover of alien fountain grass (  Pennisetum setaceum) and predation by alien rodent species. Our results indicate that although ungulate exclusion may be a necessary and critical first step, it is not sufficient to adequately preserve and maintain Hawaii's remaining tropical dry forest remnants. Our recent efforts to control the dominant alien species within the fenced preserve suggest that this practice may facilitate both the regeneration of native species and the colonization and potential invasion of new alien plants. Comparisons of seedlings of the dominant native canopy tree Diospyros sandwicensis growing in sites both dominated by and free of fountain grass suggested that fountain grass inhibits Diospyros seedling growth and photosynthesis but may increase survival if seedlings are protected from ungulates.

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Biological soil crusts (BSCs) are comprised of soil particles, bacteria, cyanobacteria, green algae, microfungi, lichens, and bryophytes and confer many ecosystem services in arid and semiarid ecosystems worldwide, including the highly threatened California sage scrub (CSS). These services, which include stabilizing the soil surface, can be adversely affected when BSCs are disturbed. Using field and greenhouse experiments, we tested the hypothesis that mechanical disturbance of BSC increases emergence of exotic vascular plants in a coastal CSS ecosystem. At Whiting Ranch Wilderness Park in southern California, 22 plots were established and emergence of exotic and native plants was compared between disturbed and undisturbed subplots containing BSC.In a separate germination study, seed fate in disturbed BSC cores was compared to seed fate in undisturbed BSC cores for three exotic and three native species. In the field, disturbed BSCs had significantly ([39) greater exotic plant emergence than in undisturbed BSC, particularly for annual grasses. Native species, however, showed no difference in emergence between disturbed and undisturbed BSC. Within the disturbed treatment, emergence of native plants was significantly, and three times less than that of exotic plants. In the germination study, seed fates for all species were significantly different between disturbed and undisturbed BSC cores. Exotic species had greater emergence in disturbed BSC, whereas native plants showed either no response or a positive response. This study demonstrates another critical ecosystem service of BSCs-the inhibition of exotic plant species-and underscores the importance of BSC conservation in this biodiversity hotspot and possibly in other aridland ecosystems.