Contrasting responses of woody and grassland ecosystems to increased CO2 as water supply varies.

Experiments show that elevated atmospheric CO2 (eCO2) often enhances plant photosynthesis and productivity, yet this effect varies substantially and may be climate sensitive. Understanding if, where and how water supply regulates CO2 enhancement is critical for projecting terrestrial responses to increasing atmospheric CO2 and climate change. Here, using data from 14 long-term ecosystem-scale CO2 experiments, we show that the eCO2 enhancement of annual aboveground net primary productivity is sensitive to annual precipitation and that this sensitivity differs between woody and grassland ecosystems. During wetter years, CO2 enhancement increases in woody ecosystems but declines in grass-dominated systems. Consistent with this difference, woody ecosystems can increase leaf area index in wetter years more effectively under eCO2 than can grassland ecosystems. Overall, and across different precipitation regimes, woody systems had markedly stronger CO2 enhancement (24%) than grasslands (13%). We developed an empirical relationship to quantify aboveground net primary productivity enhancement on the basis of changes in leaf area index, providing a new approach for evaluating eCO2 impacts on the productivity of terrestrial ecosystems.

Dominant plant taxa predict plant productivity responses to CO2 enrichment across precipitation and soil gradients.

The Earth's atmosphere will continue to be enriched with carbon dioxide (CO2) over the coming century. Carbon dioxide enrichment often reduces leaf transpiration, which in water-limited ecosystems may increase soil water content, change species abundances and increase the productivity of plant communities. The effect of increased soil water on community productivity and community change may be greater in ecosystems with lower precipitation, or on coarser-textured soils, but responses are likely absent in deserts. We tested correlations among yearly increases in soil water content, community change and community plant productivity responses to CO2 enrichment in experiments in a mesic grassland with fine- to coarse-textured soils, a semi-arid grassland and a xeric shrubland. We found no correlation between CO2-caused changes in soil water content and changes in biomass of dominant plant taxa or total community aboveground biomass in either grassland type or on any soil in the mesic grassland (P > 0.60). Instead, increases in dominant taxa biomass explained up to 85 % of the increases in total community biomass under CO2 enrichment. The effect of community change on community productivity was stronger in the semi-arid grassland than in the mesic grassland, where community biomass change on one soil was not correlated with the change in either the soil water content or the dominant taxa. No sustained increases in soil water content or community productivity and no change in dominant plant taxa occurred in the xeric shrubland. Thus, community change was a crucial driver of community productivity responses to CO2 enrichment in the grasslands, but effects of soil water change on productivity were not evident in yearly responses to CO2 enrichment. Future research is necessary to isolate and clarify the mechanisms controlling the temporal and spatial variations in the linkages among soil water, community change and plant productivity responses to CO2 enrichment.

Spatio-temporal analysis of remotely-sensed forest mortality associated with road de-icing salts.

Forest mortality along highways has long been a concern in areas where de-icing compounds are applied during winter. This study combined the spatial advantage of high-resolution remote sensing imagery and the temporal advantage of long-term archival imagery to quantify forest mortality and to detect the subtle and chronic effects of road de-icing salts for a large mountain watershed in the Sierra Nevada Mountains, USA. IKONOS-derived mortality was used in a fine-scale spatial analysis to assess road proximity and roadside topography effects on forest mortality and to compare two potential mechanisms of de-icing salt damage, i.e. aerial deposition and soil uptake. These mechanisms were modeled using spatial proxy variables that were constructed from LiDAR topographical data. The analysis revealed a clear trend of increasing mortality with increasing potential for aerial deposition of de-icing salt onto tree crowns, mainly occurring within 10 m from roads. The effect of soil uptake of salt was weaker than that of aerial deposition but had a broader potential effect zone that extended to at least 100 m from roads. Landsat TM-derived mortality from 1989 to 2010 provided a long-term time series that indicated both immediate and lagged effects of salt application on forest mortality. Immediate effects of de-icing salt were only distinct in wet years when salt application and spray generation by passing traffic and snow plowing were likely high and other damaging factors, such as bark beetles or drought mortality, were likely weak. A strong and consistent one-year lag in the effect of salt application on incidence of mortality suggested that longer-term impacts of de-icing salt on forest health likely involved more complex pathways than simply aerial deposition. Our multi-scale remote sensing approach provided convincing evidence that de-icing salt was a significant factor for roadside forest mortality and allows for efficient future monitoring at the large-watershed scale.

No cumulative effect of 10 years of elevated [CO2 ] on perennial plant biomass components in the Mojave Desert.

Elevated atmospheric CO2 concentrations ([CO2 ]) generally increase primary production of terrestrial ecosystems. Production responses to elevated [CO2 ] may be particularly large in deserts, but information on their long-term response is unknown. We evaluated the cumulative effects of elevated [CO2 ] on primary production at the Nevada Desert FACE (free-air carbon dioxide enrichment) Facility. Aboveground and belowground perennial plant biomass was harvested in an intact Mojave Desert ecosystem at the end of a 10-year elevated [CO2 ] experiment. We measured community standing biomass, biomass allocation, canopy cover, leaf area index (LAI), carbon and nitrogen content, and isotopic composition of plant tissues for five to eight dominant species. We provide the first long-term results of elevated [CO2 ] on biomass components of a desert ecosystem and offer information on understudied Mojave Desert species. In contrast to initial expectations, 10 years of elevated [CO2 ] had no significant effect on standing biomass, biomass allocation, canopy cover, and C : N ratios of above- and belowground components. However, elevated [CO2 ] increased short-term responses, including leaf water-use efficiency (WUE) as measured by carbon isotope discrimination and increased plot-level LAI. Standing biomass, biomass allocation, canopy cover, and C : N ratios of above- and belowground pools significantly differed among dominant species, but responses to elevated [CO2 ] did not vary among species, photosynthetic pathway (C3 vs. C4 ), or growth form (drought-deciduous shrub vs. evergreen shrub vs. grass). Thus, even though previous and current results occasionally show increased leaf-level photosynthetic rates, WUE, LAI, and plant growth under elevated [CO2 ] during the 10-year experiment, most responses were in wet years and did not lead to sustained increases in community biomass. We presume that the lack of sustained biomass responses to elevated [CO2 ] is explained by inter-annual differences in water availability. Therefore, the high frequency of low precipitation years may constrain cumulative biomass responses to elevated [CO2 ] in desert environments.

Do increased summer precipitation and N deposition alter fine root dynamics in a Mojave Desert ecosystem?

Climate change is expected to impact the amount and distribution of precipitation in the arid southwestern United States. In addition, nitrogen (N) deposition is increasing in these regions due to increased urbanization. Responses of belowground plant activity to increases in soil water content and N have shown inconsistent patterns between biomes. In arid lands, plant productivity is limited by water and N availability so it is expected that changes in these factors will affect fine root dynamics. The objectives of this study were to quantify the effects of increased summer precipitation and N deposition on fine root dynamics in a Mojave Desert ecosystem during a 2-year field experiment using minirhizotron measurements. Root length density, production, and mortality were measured in field plots in the Mojave Desert receiving three 25 mm summer rain events and/or 40 kg N ha(-1)  yr(-1) . Increased summer precipitation and N additions did not have an overall significant effect on any of the measured root parameters. However, differences in winter precipitation resulting from interannual variability in rainfall appeared to affect root parameters with root production and turnover increasing following a wet winter most likely due to stimulation of annual grasses. In addition, roots were distributed more deeply in the soil following the wet winter. Root length density was initially higher under canopies compared to canopy interspaces, but converged toward the end of the study. In addition, roots tended to be distributed more deeply into the soil in canopy interspace areas. Results from this study indicated that increased summer precipitation and N deposition in response to climate change and urbanization are not likely to affect fine root dynamics in these Mojave Desert ecosystems, despite studies showing aboveground plant physiological responses to these environmental perturbations. However, changes in the amount and possibly distribution of winter precipitation may affect fine root dynamics.

Temporal dynamics of fine roots under long-term exposure to elevated CO2 in the Mojave Desert.

Deserts are considered 'below-ground dominated', yet little is known about the impact of rising CO(2) in combination with natural weather cycles on long-term dynamics of root biomass. This study quantifies the temporal dynamics of fine-root production, loss and standing crop in an intact desert ecosystem exposed to 10 yr of elevated CO(2). We used monthly minirhizotron observations from 4 yr (2003-2007) for two dominant shrub species and along community transects at the Nevada Desert free-air CO(2) enrichment Facility. Data were synthesized within a Bayesian framework that included effects of CO(2) concentration, cover type, phenological period, antecedent soil water and biological inertia (i.e. the influence of prior root production and loss). Elevated CO(2) treatment interacted with antecedent soil moisture and had significantly greater effects on fine-root dynamics during certain phenological periods. With respect to biological inertia, plants under elevated CO(2) tended to initiate fine-root growth sooner and sustain growth longer, with the net effect of increasing the magnitude of production and mortality cycles. Elevated CO(2) interacts with past environmental (e.g. antecedent soil water) and biological (e.g. biological inertia) factors to affect fine-root dynamics, and such interactions are expected to be important for predicting future soil carbon pools.

Physiological and morphological responses of Tamarix ramosissima and Populus euphratica to altered groundwater availability.

Riparian plants in arid areas are subject to frequent hydrological fluctuations induced through natural flow variation and water use by humans. Although many studies have focused on the success of Tamarix ramosissima Ledeb. in its invaded ranges, its major competitor in its home range, Populus euphratica Oliv., historically has dominated riparian forests where both species occur naturally. Thus, identifying ecophysiological differences between T. ramosissima and its co-evolved competitor under varying hydrological conditions may help us understand how flow regimes affect dominance in its home range and promote invasion in new ranges. We examined ecophysiological responses of T. ramosissima and P. euphratica, which are both native to the Tarim River Basin, northwest China, to experimental alterations in groundwater. Seedlings of both species were grown in lysimeters, first under well-watered conditions and then exposed to different groundwater treatments: inundation, drought, and relatively shallow, moderate and deep groundwater. Under inundation, T. ramosissima showed little growth whereas P. euphratica died after ~45 days. Droughted seedlings of both species suffered from considerable water stress evidenced by slow growth, decreased total leaf area and specific leaf area, and decreased xylem water potential (ψ), maximum photosynthetic rate and carboxylation efficiency. Both species had better ecophysiological performances under shallow and moderate groundwater conditions. When groundwater declined below rooting depth, seedlings of both species initially experienced decreased ψ, but ψ of T. ramosissima recovered late in the experiment whereas P. euphratica maintained decreased ψ. This ability of T. ramosissima to recover from water deficit might result from its rapid root elongation and subsequent ability to acquire groundwater, which in turn likely provides ecophysiological advantages over P. euphratica. Our results suggest that recent groundwater declines along the Tarim River could facilitate T. ramosissima more due to its rapid response to changed groundwater availability. This trait may also help the success of T. ramosissima as it invaded riparian ecosystems in southwestern USA.

Maintenance of C sinks sustains enhanced C assimilation during long-term exposure to elevated [CO2] in Mojave Desert shrubs.

During the first few years of elevated atmospheric [CO(2)] treatment at the Nevada Desert FACE Facility, photosynthetic downregulation was observed in desert shrubs grown under elevated [CO(2)], especially under relatively wet environmental conditions. Nonetheless, those plants maintained increased A (sat) (photosynthetic performance at saturating light and treatment [CO(2)]) under wet conditions, but to a much lesser extent under dry conditions. To determine if plants continued to downregulate during long-term exposure to elevated [CO(2)], responses of photosynthesis to elevated [CO(2)] were examined in two dominant Mojave Desert shrubs, the evergreen Larrea tridentata and the drought-deciduous Ambrosia dumosa, during the eighth full growing season of elevated [CO(2)] treatment at the NDFF. A comprehensive suite of physiological processes were collected. Furthermore, we used C labeling of air to assess carbon allocation and partitioning as measures of C sink activity. Results show that elevated [CO(2)] enhanced photosynthetic performance and plant water status in Larrea, especially during periods of environmental stress, but not in Ambrosia. δ(13)C analyses indicate that Larrea under elevated [CO(2)] allocated a greater proportion of newly assimilated C to C sinks than Ambrosia. Maintenance by Larrea of C sinks during the dry season partially explained the reduced [CO(2)] effect on leaf carbohydrate content during summer, which in turn lessened carbohydrate build-up and feedback inhibition of photosynthesis. δ(13)C results also showed that in a year when plant growth reached the highest rates in 5 years, 4% (Larrea) and 7% (Ambrosia) of C in newly emerging organs were remobilized from C that was assimilated and stored for at least 2 years prior to the current study. Thus, after 8 years of continuous exposure to elevated [CO(2)], both desert perennials maintained their photosynthetic capacities under elevated [CO(2)]. We conclude that C storage, remobilization, and partitioning influence the responsiveness of these desert shrubs during long-term exposure to elevated [CO(2)].

Transitory effects of elevated atmospheric CO2 on fine root dynamics in an arid ecosystem do not increase long-term soil carbon input from fine root litter.

Experimental increases in atmospheric CO2 often increase root production over time, potentially increasing soil carbon (C) sequestration. Effects of elevated atmospheric CO2 on fine root dynamics in a Mojave desert ecosystem were examined for the last 4.5 yr of a long-term (10-yr) free air CO2 enrichment (FACE) study at the Nevada desert FACE facility (NDFF). Sets of minirhizotron tubes were installed at the beginning of the NDFF experiment to characterize rooting dynamics of the dominant shrub Larrea tridentata, the codominant shrub Ambrosia dumosa and the plant community as a whole. Although significant treatment effects occurred sporadically for some fine root measurements, differences were transitory and often in opposite directions during other time-periods. Nonetheless, earlier root growth under elevated CO2 helped sustain increased assimilation and shoot growth. Overall CO2 treatment effects on fine root standing crop, production, loss, turnover, persistence and depth distribution were not significant for all sampling locations. These results were similar to those that occurred near the beginning of the NDFF experiment but unlike those in other ecosystems. Thus, increased C input into soils is unlikely to occur from fine root litter under elevated atmospheric CO2 in this arid ecosystem.

Invasion triangle: an organizational framework for species invasion.

Species invasion is a complex, multifactor process. To encapsulate this complexity into an intuitively appealing, simple, and straightforward manner, we present an organizational framework in the form of an invasion triangle. The invasion triangle is an adaptation of the disease triangle used by plant pathologists to help envision and evaluate interactions among a host, a pathogen, and an environment. Our modification of this framework for invasive species incorporates the major processes that result in invasion as the three sides of the triangle: (1) attributes of the potential invader; (2) biotic characteristics of a potentially invaded site; and (3) environmental conditions of the site. The invasion triangle also includes the impact of external influences on each side of the triangle, such as climate and land use change. This paper introduces the invasion triangle, discusses how accepted invasion hypotheses are integrated in this framework, describes how the invasion triangle can be used to focus research and management, and provides examples of application. The framework provided by the invasion triangle is easy to use by both researchers and managers and also applicable at any level of data intensity, from expert opinion to highly controlled experiments. The organizational framework provided by the invasion triangle is beneficial for understanding and predicting why species are invasive in specific environments, for identifying knowledge gaps, for facilitating communication, and for directing management in regard to invasive species.

Are Mojave Desert annual species equal? Resource acquisition and allocation for the invasive grass Bromus madritensis subsp. rubens (Poaceae) and two native species.

Abundance of invasive plants is often attributed to their ability ot outcompete native species. We compared resource acquisition and allocation of the invasive annual grass Bromus madritensis subsp. rubens with that of two native Mojave Desert annuals, Vulpia octoflora and Descurainia pinnata, in a glasshouse experiment. Each species was grown in monoculture at two densities and two levels of N availability to compare how these annuals capture resources and to understand their relative sensitivities to environmental change. During >4 mo of growth, Bromus used water more rapidly and had greater biomass and N content than the natives, partly because of its greater root-surface area and its exploitation of deep soils. Bromus also had greater N uptake, net assimilation and transpiration rates, and canopy area than Vulpia. Resource use by Bromus was less sensitive to changes in N availability or density than were the natives. The two native species in this study produced numerous small seeds that tended to remain dormant, thus ensuring escape of offspring from unfavorable germination conditions; Bromus produced fewer but larger seeds that readily germinated. Collectively, these traits give Bromus the potential to rapidly establish in diverse habitats of the Mojave Desert, thereby gaining an advantage over coexisting native species.

Gas exchange and carbon isotope discrimination of Juniperus osteosperma and Juniperus occidentalis across environmental gradients in the Great Basin of western North America.

We determined how ecophysiological characteristics of two juniper species, Juniperus occidentalis Hook. (western juniper) and Juniperus osteosperma (Torr.) Little (Utah juniper), changed along altitudinal and regional environmental gradients in the Great Basin of western North America. We obtained diurnal measurements of leaf gas exchange and xylem water potential (Psi) from plants at a low and a high altitude site within each of six mountain ranges during fall 1994, spring, summer, and fall 1995, and summer 1996. We also determined carbon isotope composition (delta(13)C) of leaf cellulose produced during the 1995 growing season. Overall, leaf gas exchange, Psi and delta(13)C did not differ significantly between species. Differences in daily (A(d)) and season-long (A(s)) carbon assimilation among mountain ranges suggested two groupings-a group of northern ranges and a group of southern ranges. Each group contained one mountain range with J. occidentalis and two with J. osteosperma. Differences in carbon assimilation based on this grouping were associated with two findings: (1) conductance of CO(2) from substomatal cavities to the site of carboxylation (g(m)) for junipers in the northern ranges averaged almost twice that of junipers in the southern ranges; and (2) physiological shifts occurred such that A(d) of junipers in the northern ranges was influenced more by Psi(pd), whereas A(d) of junipers in the southern ranges was influenced more by leaf temperature. Mean delta(13)C over all trees at a site was significantly correlated with annual precipitation. Significant differences in A(d) occurred between altitudes, but these differences were associated with differences in the timing of optimum leaf temperature for photosynthesis rather than with physiological acclimation to temperature, irradiance, or Psi. Most gas exchange parameters (e.g., assimilation, transpiration, stomatal conductance, and water use efficiency) varied seasonally, and the seasonal differences were strongly influenced by water stress.

Soil water exploitation after fire: competition between Bromus tectorum (cheatgrass) and two native species.

Causes for the widespread abundance of the alien grass Bromus tectorum (cheatgrass) after fire in semiarid areas of western North America may include: (1) utilization of resources freed by the removal of fireintolerant plants; and (2) successful competition between B. tectorum and individual plants that survive fire. On a site in northwestern Nevada (USA), measurements of soil water content, plant water potential, aboveground biomass production, water use efficiency, and B. tectorum tiller density were used to determine if B. tectorum competes with either of two native species (Stipa comata and Chrysothamnus viscidiflorus) or simply uses unclaimed resources. Soil water content around native species occurring with B. tectorum was significantly lower (P<0.05) than around individuals without B. tectorum nearby. Native species had significantly more negative plant water potential when they occurred with B. tectorum. Aboveground biomass was significantly higher for native species without B. tectorum. However, the carbon isotope ratio of leaves for native species with B. tectorum was not significantly different from individuals without B. tectorum. Thus, B. tectorum competes with native species for soil water and negatively affects their wate status and productivity, but the competition for water does not affect water use efficiency of the native species. These adverse effects of B. tectorum competition on the productivity and water status of native species are also evident at 12 years after a fire. This competitive ability of B. tectorum greatly enhances its capability to exploit soil resources after fire and to enhance its status in the community.

Gas exchange of Agropyron desertorum: diurnal patterns and responses to water vapor gradient and temperature.

The response of leaf gas exchange to environmental variables were measured at different levels of drought stress for Agropyron desertorum, a naturalized perennial bunchgrass of the semiarid shrub steppes of western North America. Leaf conductance (stomatal plus boundary layer) was more sensitive to changes in water vapor gradient than to changes in leaf temperature. Assimilation was sensitive to both temperature and vapor gradient, and also appeared to be affected by conductance and high transpiration rates. The magnitudes of both assimilation and conductance decreased with increased drought conditions. Diurnal patterns of gas exchange were measured during 3 growing seasons. For a typical spring day with moderate leaf temperature and vapor gradient, diurnal patterns were similar for plants at different levels of soil water availability. Assimilation was relatively constant during most of the day, but conductance decreased during the afternoon. Total daily carbon gain was decreased to a lesser extent than daily water loss as soil water was depleted. Consequently, the ratio of daily carbon gain to daily water loss, i.e. daily water use efficiency, increased with decreased soil water content for diurnals under spring conditions. Diurnal patterns of assimilation for a typical summer day with high leaf temperature and vapor gradient differend from those for a spring day. An afternoon decrease in assimilation was typical during a summer day. Daily carbon gain, water use, and water use efficiency for summer diurnals decreased only under severe drought conditions. Almost complete recovery of assimilation and conductance occurred if leaf microclimate was ameliorated during the afternoon of either spring or summer diurnals. Thus, conditions responsible for a midday depression in assimilation during a single day did not have persistent effects on leaf gas exchange. Daily carbon gain of a typical summer day was restricted by leaf microclimate during the afternoon, but daily water use efficiency was not relatively increased by the amelioration of leaf microclimate.