A. Joshua Leffler

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.

HYDRAULIC REDISTRIBUTION THROUGH THE ROOT SYSTEMS OF SENESCED PLANTS

Hydraulic redistribution, the movement of water from soil layers of higher water potential to layers of lower water potential through the root systems of plants, has been documented in many taxa worldwide. Hydraulic redistribution is influenced principally by physical properties of roots and soils, and it should occur whenever root systems span soil layers of different water potential. Therefore, hydraulic redistribution should occur through the root systems of plants with aboveground tissue removed or through the root systems of fully senesced plants as long as roots remain intact and hydrated. We examined our hypothesis in field and greenhouse studies with the annual grass Bromus tectorum. We used soil psychrometry to measure soil water potential and performed 2H-labeling exper- iments. In the field, following senescence of B. tectorum, we show substantial changes in soil water potential consistent with both upward and downward movement of water through roots. The amount of water redistributed represents a significant proportion of that which can be stored in the rooted zone. We also experimentally demonstrated upward movement of a 2H label by roots of senesced plants and by roots of plants without aboveground tissues. In the greenhouse, we further demonstrated redistribution by senesced individuals using a 2H label. Hydraulic redistribution through the roots of senesced plants should receive further attention because it may have important ecological consequences for soil water recharge, survival of plants through drought, and agricultural practices.

Long-term tree ring chronologies from sympatric tropical dry-forest trees: individualistic responses to climatic variation

The influence of local precipitation and temperature on long-term growth dynamics in two species of seasonally dry tropical forest trees were investig- ated. Growth records were extracted from tree rings in Guanacaste province, Costa Rica. These chronologies provide a long-term (c. 85-y) record of tree growth for two species with contrasting phenologies. Annual growth, in both species, was dependent on annual and/or monthly variation in local precipitation but less so on temperature. For each species, however, patterns of growth reflected unique degrees of sensitivity to monthly rainfall and rainfall during previous years. It is hypothesized that such differences were due to the rooting depth of these species. A review of the literature also indicated similar diverse cambial growth responses by tropical trees to variation in annual and monthly climate. Lastly, it was shown that variation in longer term fluctuations in the Pacific and Atlantic oceans, as measured by the El Nifio Southern Oscillation (ENSO) and North Atlantic Oscilla- tion (NAO), significantly influenced local precipitation in Guanacaste only during the wettest portion of the wet season. Such temporal sensitivity may have differen- tially influenced the longer-term growth of some tropical tree species but not others. Together, these results support the hypothesis that tropical tree species respond individualistically to variation in local and regional climate and that some tropical assemblages may in fact be structured by species-specific differences in soil water-use.

Functional Differences in Soil Water Pools: a New Perspective on Plant Water Use in Water-Limited Ecosystems

Arid and semi-arid ecosystems cover roughly half of the earths surface. Significant changes in vegetation cover combined with climate change have increased concern over the future of these lands, which have considerable economic importance. Much research has focused on plant-soil water relations in these systems, yet many mechanisms and significance of water use patterns are not well under- stood. Here we describe a new conceptual model that considers two pools of soil water accessed by plants: a growth pool that is located in shallow soil layers, and a maintenance pool that is often in deeper soil layers. While they may be spatially and

Response of Water Vapor and CO2 Fluxes in Semiarid Lands to Seasonal and Intermittent Precipitation Pulses

Abstract Precipitation pulses are important in controlling ecological processes in semiarid ecosystems. The effects of seasonal and intermittent precipitation events on net water vapor and CO2 fluxes were determined for crested wheatgrass (Agropyron desertorum), juniper (Juniperus osteosperma), and sagebrush (Artemisia tridentata) ecosystems using eddy covariance measurements. The measurements were made at Rush Valley, Utah, in the northern Great Basin of the United States. Data were evaluated during the growing seasons of 2002 and 2003. Each of these communities responds to precipitation pulses in all seasons, but these responses vary among season and ecosystem, and differ for water vapor and CO2. The degree and direction of response (i.e., net uptake or efflux) depended upon the timing and amount of precipitation. In early spring, both evapotranspiration (ET) and CO2 fluxes responded only slightly to precipitation pulses because soils were already moist from snowmelt and spring rains. As soils dried lat...

A new perspective on trait differences between native and invasive exotic plants

Functional differences between native and exotic species potentially constitute one factor responsible for plant invasion. Differences in trait values between native and exotic invasive species, however, should not be considered fixed and may depend on the context of the comparison. Furthermore, the magnitude of difference between native and exotic species necessary to trigger invasion is unknown. We propose a criterion that differences in trait values between a native and exotic invasive species must be greater than differences between co-occurring natives for this difference to be ecologically meaningful and a contributing factor to plant invasion. We used a meta-analysis to quantify the difference between native and exotic invasive species for various traits examined in previous studies and compared this value to differences among native species reported in the same studies. The effect size between native and exotic invasive species was similar to the effect size between co-occurring natives except for studies conducted in the field; in most instances, our criterion was not met although overall differences between native and exotic invasive species were slightly larger than differences between natives. Consequently, trait differences may be important in certain contexts, but other mechanisms of invasion are likely more important in most cases. We suggest that using trait values as predictors of invasion will be challenging.

Gas exchange and growth responses of the desert shrubs Artemisia tridentata and Chrysothamnus nauseosus to shallow- vs. deep-soil water in a glasshouse experiment

Abstract The aridland shrub species, Artemisia tridentata (big sagebrush) and Chrysothamnus nauseosus (rubber rabbitbrush), are distributed widely in the Intermountain region of western North America. Earlier research indicated that A. tridentata can utilize upper soil water from transient summer rain events while C. nauseosus apparently cannot, although both species have similar rooting depths. Thus, we hypothesized that C. nauseosus relies more on deep water than A. tridentata , while A. tridentata can take advantage of soil moisture in upper soil layers. We examined this hypothesis by growing A. tridentata and C. nauseosus in two-layer pots in which soil water content in the upper and lower layers was controlled independently. After plants were well established, they were subjected to one of the three water treatments: water applied both to upper and lower layers, water applied only to the upper layer, or water applied only to the lower layer. We measured above- and belowground biomass, leaf gas exchange and leaf carbon isotope composition (δ 13 C). Close to the end of the experiment, a deuterium- and 15 N-labeled solution was applied in a localized patch in either the upper or lower soil layer to quantify resource-pulse utilization. In general, our hypothesis was supported; δ 13 C and biomass indicated that A. tridentata performed better when water was available in the upper soil layer and gas exchange, and δ 13 C indicated that C. nauseosus performed better when water was available in the lower layer. There was, however, no significant variation among treatments for many of the variables examined. Roots of both species were involved in hydraulic redistribution of soil water between layers, which may have reduced the effectiveness of the water distribution treatments to some extent.

Carbon isotope composition of tree leaves from Guanacaste, Costa Rica: comparison across tropical forests and tree life history

Despite the progress made in understanding the ecophysiology of tropical plants during the past two decades (Luttge 1997, Mulkey et al. 1996), questions regarding relationships between the environment and physiological diversity remain. It is now recognized that tropical climate can be quite variable (see Coen 1983) which could lead to significant functional diversity (increased variation in life history traits) among species due to the tight association between gas exchange physiology and the environment (see Enquist & Leffler 2001, Guehl et al. 1998, Huc et al. 1994, Martinelli et al. 1998, Sobrado 1993). It remains unclear, however, how the subtleties of variation in tropical climate and tree life history traits are related to the functional diversity of tropical communities (Borchert 1994, 1998).

Invasion is Contingent on Species Assemblage and Invasive Species Identity in Experimental Rehabilitation Plots

Abstract Ecological studies often suggest that diverse communities are most resistant to invasion by exotic plants, but relatively few local species may be available to a rehabilitation practitioner. We examine the ability of monocultures and diverse assemblages to resist invasion by an exotic annual grass (cheatgrass) and an exotic biennial forb (dyers woad) in experimental rehabilitation plots. We constructed seven assemblages that included three monocultures of grass, forb, or shrub; three four-species mixtures of grasses, forbs, or shrubs; and a three-species mixture of one species from each growth form in an experimental field setting to test resistance to invasion. Assemblages were seeded with cheatgrass and dyers woad for two consecutive years and quantified as biomass and density of individuals from each exotic species. Soil NO3− and leaf-area index were examined as predictors of invasive plant abundance. Cheatgrass invasion was greatest in forb and shrub assemblages, and least in mixed grass or grass monoculture; dyers woad invasion was greatest into mixed grass or grass monoculture, but least into monoculture or mixed-species assemblages composed of forbs or shrubs. The community composed of grasses, forbs, and shrubs suppressed invasion by both species. Consequently, assemblages were most resistant to invasion by species of the same growth form. Moreover, these monocultures and mixtures were generally similar in conferring resistance to invasion, but a monoculture of big sagebrush was more resistant than a mixture of shrubs. Soil NO3− was correlated with invasion by cheatgrass, whereas LAI was correlated with invasion by dyers woad, suggesting these species were more limited by belowground and aboveground resources, respectively. Overall, increasing diversity with limited species did not necessarily enhance resistance to invasion.

Thermal stress response in a dinoflagellate-bearing nudibranch and the octocoral on which it feeds

Abstract In this study, we examined two non-scleractinian taxa, the rare nudibranch Phyllodesmium lizardensis and Bayerxenia sp., the octocoral on which the nudibranch lives and feeds, to investigate the effect of experimental heat stress on their symbioses with Symbiodinium. Bleaching has not been studied in nudibranchs. Bayerxenia sp. belongs to the alcyonacea family Xeniidae, members of which are known to be heat sensitive, but the genus has never been subject to heat stress experiments or bleaching observations. While qPCR did not reveal any changes to the symbiont community composition, the two host species responded differently to increased temperature. There were changes in the relative proportion of tissue types in Bayerxenia sp., but these were not attributable to the temperature treatment. Bayerxenia sp. exhibited no changes in cellular structure (apoptosis or cell necrosis), or symbiont functioning, cell size, density, or cladal community structure. On the other hand, the host, P. lizardensis, experienced tissue loss and symbiont densities decreased significantly with the majority of the remaining symbiont cells significantly degenerated after the heat stress. This decrease did not influence symbiont community composition, symbiont cell size, or photosynthetic efficiency. While the bleaching process in nudibranchs was demonstrated for the first time, the physiological and molecular pathways leading to this response still require attention.