Global-change drivers of ecosystem functioning modulated by natural variability and saturating responses.

Humans are altering global environment at an unprecedented rate through changes in biodiversity, climate, nitrogen cycle, and land use. To address their effects on ecosystem functioning, experiments most frequently explore one driver at a time and control as many confounding factors as possible. Yet, which driver exerts the largest influence on ecosystem functioning and whether their relative importance changes among systems remain unclear. We analyzed experiments in the Patagonian steppe that evaluated the aboveground net primary production (ANPP) response to manipulated gradients of species richness, precipitation, temperature, nitrogen fertilization (N), and grazing intensity. We compared the effect on ANPP relative to ambient conditions considering intensity and direction of manipulations for each driver. The ranking of responses to drivers with comparable manipulation intensity was as follows: biodiversity>grazing>precipitation>N. For a similar intensity of manipulation, the effect of biodiversity loss was 4.0, 3.6, and 1.5, times larger than N deposition, decreased precipitation, and increased grazing intensity. We interpreted our results considering two hypotheses. First, the response of ANPP to changes in precipitation and biodiversity is saturating, so we expected larger effects when the driver was reduced, relative to ambient conditions, than when it was increased. Experimental manipulations that reduced ambient levels had larger effects than those that increased them. Second, the sensitivity of ANPP to each driver is inversely related to the natural variability of the driver. In Patagonia, the ranking of natural variability of drivers is as follows: precipitation>grazing>temperature>biodiversity>N. So, in general, the ecosystem was most sensitive to drivers that varied the least. Comparable results from Cedar Creek (MN) support both hypotheses and suggest that sensitivity to drivers varies among ecosystem types. Given the importance of understanding ecosystem sensitivity to predict global-change impacts, it is necessary to design new experiments located in regions with contrasting natural variability and that include the full range of drivers.

Effect of woody-plant encroachment on livestock production in North and South America.

A large fraction of the world grasslands and savannas are undergoing a rapid shift from herbaceous to woody-plant dominance. This land-cover change is expected to lead to a loss in livestock production (LP), but the impacts of woody-plant encroachment on this crucial ecosystem service have not been assessed. We evaluate how tree cover (TC) has affected LP at large spatial scales in rangelands of contrasting social-economic characteristics in the United States and Argentina. Our models indicate that in areas of high productivity, a 1% increase in TC results in a reduction in LP ranging from 0.6 to 1.6 reproductive cows (Rc) per km(2). Mean LP in the United States is 27 Rc per km(2), so a 1% increase in TC results in a 2.5% decrease in mean LP. This effect is large considering that woody-plant cover has been described as increasing at 0.5% to 2% per y. On the contrary, in areas of low productivity, increased TC had a positive effect on LP. Our results also show that ecological factors account for a larger fraction of LP variability in Argentinean than in US rangelands. Differences in the relative importance of ecological versus nonecological drivers of LP in Argentina and the United States suggest that the valuation of ecosystem services between these two rangelands might be different. Current management strategies in Argentina are likely designed to maximize LP for various reasons we are unable to explore in this effort, whereas land managers in the United States may be optimizing multiple ecosystem services, including conservation or recreation, alongside LP.

Interactions among resource partitioning, sampling effect, and facilitation on the biodiversity effect: a modeling approach.

Resource partitioning, facilitation, and sampling effect are the three mechanisms behind the biodiversity effect, which is depicted usually as the effect of plant-species richness on aboveground net primary production. These mechanisms operate simultaneously but their relative importance and interactions are difficult to unravel experimentally. Thus, niche differentiation and facilitation have been lumped together and separated from the sampling effect. Here, we propose three hypotheses about interactions among the three mechanisms and test them using a simulation model. The model simulated water movement through soil and vegetation, and net primary production mimicking the Patagonian steppe. Using the model, we created grass and shrub monocultures and mixtures, controlled root overlap and grass water-use efficiency (WUE) to simulate gradients of biodiversity, resource partitioning and facilitation. The presence of shrubs facilitated grass growth by increasing its WUE and in turn increased the sampling effect, whereas root overlap (resource partitioning) had, on average, no effect on sampling effect. Interestingly, resource partitioning and facilitation interacted so the effect of facilitation on sampling effect decreased as resource partitioning increased. Sampling effect was enhanced by the difference between the two functional groups in their efficiency in using resources. Morphological and physiological differences make one group outperform the other; once these differences were established further differences did not enhance the sampling effect. In addition, grass WUE and root overlap positively influence the biodiversity effect but showed no interactions.

Preference for different inorganic nitrogen forms among plant functional types and species of the Patagonian steppe.

We have explored species-specific preferences for nitrate (NO3(-)) and ammonium (NH4(+)) as an alternative niche separation in ecosystems where nitrogen (N) is present mostly in inorganic forms. The Patagonian steppe is dominated by shrubs and grasses. Shrubs absorb water and nutrients from deep soil layers, which are poor in N, while grasses have the opposite pattern, absorbing most of their water and nutrients from the upper layers of the soil. We hypothesized that the preferences of shrub and grass for inorganic N forms are different and that the rate of potential N uptake is greater in shrubs than in grasses. To test this hypothesis, we grew individuals of six dominant species in solutions of different NH4(+):NO3(-) concentration ratios. Nitrate uptake was found to be higher in shrubs, while ammonium uptake was similar between plant functional types. The NH4(+):NO3(-) uptake ratio was significantly lower for shrubs than grasses. Shrubs, which under field conditions have deeper rooting systems than grasses, showed a higher N absorption capacity than grasses and a preference for the more mobile N form, nitrate. Grasses, which had lower N uptake rates than shrubs, preferred ammonium over nitrate. These complementary patterns between grasses and shrubs suggest a more thorough exploitation of resources by diverse ecosystems than those dominated by just one functional type. The loss of one plant functional group or a significant change in its abundance would therefore represent a reduction in resource use efficiency and ecosystem functioning.

Effects of plant species traits on ecosystem processes: experiments in the Patagonian steppe.

Several experiments have shown that aboveground net primary productivity increases with plant species richness. The main mechanism proposed to explain this relationship is niche complementarity, which is determined by differences in plant traits that affect resource use. We combined field and laboratory experiments using the most abundant species of the Patagonian steppe to identify which are the traits that determine niche complementarity in this ecosystem. We estimated traits that affect carbon, water, microclimate, and nitrogen dynamics. The most important traits distinguishing among species, from the standpoint of their effects on ecosystem functioning, were potential soil nitrification, rooting depth, and soil thermal amplitude. Additionally, we explored the relationship between trait diversity and aboveground net primary production (ANPP) using a manipulative field experiment. ANPP and the fraction of ANPP accounted for by trait diversity increased with number of traits. The effect of trait diversity decreased as the number of traits increased. Here, the use of traits gave us a mechanistic understanding of niche complementarity in the Patagonian steppe.

Vegetation structure constrains primary production response to water availability in the Patagonian steppe.

Grassland aboveground net primary production (ANPP) increases linearly with precipitation in space and time, but temporal models relating time series of ANPP and annual precipitation for single sites show lower slopes and regression coefficients than are shown by spatial models. The analysis of several ANPP time series showed lags in the ecosystem response to increased water availability, which may explain the difference between spatial and temporal models. The lags may result from constraints that ecosystems experience after drought. Our objective was to explore the structural constraints of the ANPP response to rainfall variability in a semiarid ecosystem, the Patagonian steppe, in southern Argentina. We designed a 3-yr rainfall manipulation experiment where we decreased water input with rainout shelters during two consecutive years, which included three levels of rainfall interception (30%, 55%, and 80%) and a control. In the third year, we irrigated one-half of the plots of each rainfall-interception treatment. We evaluated the immediate effects of drought on current-year ANPP and the effects of previous-year drought on vegetation recovery after water supplementation. ANPP (g x m(-2) x yr(-1)) was linearly related to annual precipitation input (APPT; mm/yr) along the experimental precipitation gradient (ANPP = 0.13 x APPT + 58.3; r2 = 0.34, P < 0.01), and this relationship was mostly accounted for by changes in the ANPP of grasses. Plant density (D; no. individuals/mm2) was related to the precipitation received during the drought period (D = 0.11 x APPT + 18; r2 = 0.39, P < 0.05). The recovery of plants after irrigation was lower for those plots that had experienced experimental drought the previous years relative to controls, and the lags were proportional to the intensity of drought. Therefore, our results suggest that the density of plants may constrain the recovery of vegetation after drought, and these constraints may determine lags that limit the capacity of the ecosystem to take advantage of wet years after dry years.

A rainout shelter design for intercepting different amounts of rainfall.

Field manipulative experiments represent a straightforward way to explore temporal relationships between annual precipitation and productivity. Water exclusion usually involves the use of rainout shelters, which are in general formed by a complete roof that intercepts 100% of the rainfall and require complicated mechanisms to move the shelter into place. The rainout-shelter design described here is a fixed-location shelter with a roof consisting of bands of transparent acrylic that blocks different amounts of rainfall while minimally affecting other environmental variables. We constructed thirty 3.76-m2 shelters in an arid steppe near Río Mayo, Argentina (at 45°41'S, 70°16'W), to impose 30%, 55%, and 80% of rainfall interception. We tested the effectiveness of the design by collecting all the intercepted water in storage tanks and we evaluated changes in soil water content with the time domain reflectometry technique. We also measured soil water content in regular grids to assess the magnitude of the edge effect. We analysed the microclimate impact of the shelters by measuring photosynthetically active radiation and air and soil temperature inside and outside shelters. We did not detect significant differences between the observed and the expected rainfall interception for the 30% and 55% interception treatments but the 80% shelters intercepted 71% of incoming rainfall, which was significantly (P<0.05) lower than the expected value. Soil water content was significantly (P<0.05) higher in the control plots than in the plots with rainout shelter at all dates, except in January (summer). Radiation was not affected by the 30% interception treatment, but the roof with the largest number of acrylics bands (80% interception treatment) reduced incident radiation throughout the day by 10%. Air and soil temperatures were lower under than outside the shelters during the period of highest radiation but the opposite occurred with low radiation but with smaller differences. The two characteristics of the shelter, fixed design and low cost, allow for proper replication in space, which is required in ecological field experiments.

Vegetación y la depresión del Salado (provincia de Buenos Aires) y su modificación por influencia antrópica

El área denominada depresión del Salado abarca una superficie aproximada de 5.800.000 ha y alrededor de un 70 por ciento de esta superficie está cubierta por pastizales naturales o modificados. El conocimiento alcanzado por el estudio de la región permitió la elección de lugares apropiados para la realización de estudios más intensivos, los cuales muestran relaciones de ciertos factores del ambiente con la presencia de algunas comunidades vegetales. Otro aspecto importante es el que se refiere a las modificaciones que el uso antrópico ha introducido en los sistemas ecológicos de la región. El sobrepastoreo y las prácticas agrícolas, aplicados en algunos de los ambientes de la región, han determinado serios cambios en ciertos caracteres de la vegetación y del suelo