Martín Oesterheld

Rooting depth, water availability, and vegetation cover along an aridity gradient in Patagonia

Above-and belowground biomass distribution, isotopic composition of soil and xylem water, and carbon isotope ratios were studied along an aridity gradient in Patagonia (44–45°S). Sites, ranging from those with Nothofagus forest with high annual rainfall (770 mm) to Nothofagus scrub (520 mm), Festuca (290 mm) and Stipa (160 mm) grasslands and into desert vegetation (125 mm), were chosen to test whether rooting depth compensates for low rainfall. Along this gradient, both mean above-and belowground biomass and leaf area index decreased, but average carbon isotope ratios of sun leaves remained constant (at-27‰), indicating no major differences in the ratio of assimilation to stomatal conductance at the time of leaf growth. The depth of the soil horizon that contained 90% of the root biomass was similar for forests and grasslands (about 0.80–0.50 m), but was shallower in the desert (0.30 m). In all habitats, roots reached water-saturated soils or ground water at 2–3 m depth. The depth profile of oxygen and hydrogen isotope ratios of soil water corresponded inversely to volumetric soil water contents and showed distinct patterns throughout the soil profile due to evaporation, water uptake and rainfall events of the past year. The isotope ratios of soil water indicated that high soil moisture at 2–3 m soil depth had originated from rainy periods earlier in the season or even from past rainy seasons. Hydrogen and oxygen isotope ratios of xylem water revealed that all plants used water from recent rain events in the topsoil and not from water-saturated soils at greater depth. However, this study cannot explain the vegetation zonation along the transect on the basis of water supply to the existing plant cover. Although water was accessible to roots in deeper soil layers in all habitats, as demonstrated by high soil moisture, earlier rain events were not fully utilized by the current plant cover during summer drought. The role of seedling establishment in determining species composition and vegetation type, and the indirect effect of seedling establishment on the use of water by fully developed plant cover, are discussed in relation to climate change and vegetation modelling.

Effect of stress and time for recovery on the amount of compensatory growth after grazing

SummaryWe tested the hypothesis that the amount of compensatory growth after defoliation is affected by the level of stress at which plants grow when defoliated and by the length of time for recovery. Growth response to defoliation went from partial compensation when plants were growing at high relative growth rates (RGR) to overcompensation when plants were more stressed and growing at low RGR. Defoliation released plants from the limitation imposed by the accumulation of old and dead tissue and this release overrode the negative effect of biomass loss. Compensatory growth resulted from a higher RGR aboveground that was not associated with a reduction in RGR belowground. Time available for recovery had a major impact on the outcome of defoliation. With a short time for recovery, RGR was decreased by defoliation because an immediate increase in net assimilation rate was overridden by a reduction in the ratio of leaf area to plant weight. After defoliation, this ratio increased quickly due to a larger allocation to leaf growth and lower leaf specific weights, resulting in higher RGR. We conclude that the compensatory response to grazing depends on the type and level of stress limiting growth. Allocation and physiological responses to stress may positively or negatively affect the response to grazing and, simultaneously, grazing may alleviate or aggravate the effects of different types of stress.

Grazing effects upon plant community structure in subhumid grasslands of Argentina

Changes in plant community structure are identified as a result of grazing in grasslands of the flooding pampa which evolved under supposedly light grazing conditions. The effect of excluding grazing upon total leaf area index was an increase of 30%. The largest response was observed in the distribution of leaves in the canopy. In the grazed areas, most of the green material was concentrated in the 0–5 cm layer while in the ungrazed treatments the largest portion of the leaf area was in the 10–30 cm layer. Grazing exclusion resulted in a small change in total basal area but a larger change in its distribution, from many small tussocks to less numerous large ones. The effect of grazing upon leaf area and basal area was accounted for by changes in vigor as well as by changes in species composition. The major effect of excluding grazing upon species composition was the disappearance of some native planophile species and most of the exotics. The species composition of grazed areas of both communities was very similar while there were large differences between the ungrazed areas and between the grazed and ungrazed areas of the same community. It is suggested that there is a group of species which responds to the coarse-grained ‘signal’ of grazing and its presence can cause dissimilar communities to converge under grazing conditions. The other group of species responded to the fine-grained ‘signal’ of the environmental conditions associated with topography.

Pathways of Grazing Effects on Soil Organic Carbon and Nitrogen

Abstract Grazing modifies the structure and function of ecosystems, affecting soil organic carbon (SOC) storage. Although grazing effects on some ecosystem attributes have been thoroughly reviewed, current literature on grazing effects on SOC needs to be synthesized. Our objective was to synthesize the effects of grazing on SOC stocks in grasslands, establishing the major mechanistic pathways involved. Additionally, and because of its importance for carbon (C) biogeochemistry, we discuss the controls of soil organic nitrogen (N) stocks. We reviewed articles analyzing grazing effects on soil organic matter (SOM) stocks by comparing grazed vs. ungrazed sites, including 67 paired comparisons. SOC increased, decreased, or remained unchanged under contrasting grazing conditions across temperature and precipitation gradients, which suggests that grazing influences the factors that control SOC accumulation in a complex way. However, our review also revealed some general patterns such as 1) root contents (a primary control of SOC formation) were higher in grazed than in their ungrazed counterparts at the driest and wettest sites, but were lower at sites with intermediate precipitation (∼400 mm to 850 mm); 2) SOM C∶N ratios frequently increased under grazing conditions, which suggests potential N limitations for SOM formation under grazing; and 3) bulk density either increased or did not change in grazed sites. Nearly all sites located in the intermediate precipitation range showed decreases or no changes in SOC. We grouped previously proposed mechanisms of grazing control over SOC into three major pathways that can operate simultaneously: 1) changes in net primary production (NPP pathway), 2) changes in nitrogen stocks (nitrogen pathway), and 3) changes in organic matter decomposition (decomposition pathway). The relative importance of the three pathways may generate variable responses of SOC to grazing. Our conceptual model suggests that rangeland productivity and soil carbon sequestration can be simultaneously increased by management practices aimed at increasing N retention at the landscape level.

Intraspecific variation in the response of Themeda triandra to defoliation: the effect of time of recovery and growth rates on compensatory growth

SummaryThe response to a single defoliation was studied on three clones of Themeda triandra collected in the short, mid, and tall grassland regions of the Serengeti National Park (Tanzania). These sites represent a gradient of decreasing grazing intensity. Growth, allocation pattern, and several morphometric traits were monitored during an 80-day period. Clipped plants of the short and medium clones fully compensated for the reduction of biomass, while plants of the tall clone showed overcompensation. During the first two weeks after clipping, clipped plants showed lower relative growth rates than unclipped ones, whereas the opposite was observed later on. Clipped plants compensated for the removal of leaf area by producing new leaves with lower specific weights and higher nitrogen content. They also produced more, smaller tillers. Although clipped plants mobilized nonstructural carbohydrates from roots and crowns, this did not account for a significant amount of growth. Relative growth rates of unclipped plants of the short clone were higher. The relative growth rate of the short clone diminished less after clipping, but also exhibited the lowest increase later. The tall clone was the most negatively affected early, but showed the highest compensation later. Compared to the other clones, the short ecotype showed many of the characteristics that defoliation induced in each individual of any clone: higher allocation to leaf area production, higher relative growth rate, higher number but smaller size of tillers, and lower leaf specific weights.

Effects of grazing on seedling establishment: the role of seed and safe-site availability

The fIrSt objective of this paper was to assess the effects of grazing on seedling establishment of two species whose relative abundance at the adult stage is affected by grazing in a contrasting fashion. Second, we evaluated the relative importance of seed versus safe-site availability in explaining the effect of grazing on seedling establishment We monitored seedling establishment on a grazed area, on two areas which had not been grazed for two and seven years, and on plots which had been exper- imentally defoliated. The species compared were Dan- thonia montevidensis, a native perennial grass which dominates both grazed and ungrazed communities, and Leontodon taraxacoides, an invading exotic rosette spe- cies from the Compositae family. Continuous grazing enhanced seedling establishment of both species through its effect on the availability of safe sites. Seed availability accounted for only one, but very important, grazing effect: the lack of response by L. taraxacoides to the defoliation in the seven-year old exclosure. Its seed supply was depleted by exclusion of grazing and, consequently, its short-tenn regeneration caDacity after disturbance was lost.

Relationship between Productivity, and Species and Functional Group Diversity in Grazed and Non-Grazed Pampas Grassland

Most hypotheses addressing the effect of diversity on ecosystem function indicate the occurrence of higher process rates with increasing diversity, and only diverge in the shape of the function depending on their assumptions about the role of individual species and functional groups. Contrarily to these predictions, we show that grazing of the Flooding Pampas grasslands increased species richness, but drastically reduced above ground net primary production, even when communities with similar initial biomass were compared. Grazing increased species richness through the addition of a number of exotic forbs, without reducing the richness and cover of the native flora. Since these forbs were essentially cool-season species, and also because their introduction has led to the displacement of warm-season grasses from dominant to subordinate positions in the community, grazing not only decreased productivity, but also shifted its seasonality towards the cool season. These results suggest that species diversity and/or richness alone are poor predictors of above-ground primary production. Therefore, models that relate productivity to diversity should take into account the relative abundance and identity of species that are added or deleted by the specific disturbances that modify diversity.

Seasonal Variation in Aboveground Production and Radiation-use Efficiency of Temperate rangelands Estimated through Remote Sensing

Aboveground net primary production (ANPP) of grasslands varies spatially and temporally. Spectral information provided by remote sensors is a promising new tool that may be able to estimate ANPP in real time and at low cost. The objectives of this study were (a) to evaluate at a seasonal scale the relationship between ANPP and the normalized difference vegetation index (NDVI), (b) to estimate seasonal variations in the coefficient of conversion of absorbed radiation into aboveground biomass (εa), and (c) to identify the environmental controls on such temporal changes. We used biomass-based field determinations of ANPP for two grassland sites in the Flooding Pampa, Argentina, and related them with NDVI data derived from the NOAA Advanced Very High Resolution Radiometer (AVHRR) satellites using three different models. Results were compared with data obtained from the new Moderate Resolution Imaging Spectroradiometer (MODIS) sensor at an additional site. The first model was based solely on NDVI; the second was based on the amount of photosynthetically active radiation absorbed by the green vegetation (APARg), which was derived from NDVI and incoming photosynthetically active radiation (PAR); the third was based on APARg and εa, which was in turn estimated from climatic variables. NDVI explained between 63 and 93% of ANPP variation, depending on the site considered. Estimates of ANPP were not improved by considering the variation in incoming PAR. At both sites, εa varied seasonally (from 0.2 to 1.2 g DM/MJ) and was significantly associated with combinations of precipitation and temperature. Combining εa variations with APARg increased our ability to account for seasonal ANPP variations at both sites. Our results indicate that NDVI produces good, direct estimates of ANPP only if NDVI, PAR, and εa are correlated throughout the seasons. Thus, in most cases, seasonal variations of εa associated with temperature and precipitation must be taken into account to generate seasonal ANPP estimates with acceptable accuracy.

RELATION BETWEEN NOAA-AVHRR SATELLITE DATA AND STOCKING RATE OF RANGELANDS

Biomass of both wild herbivores and livestock in rangelands is correlated with rainfall at a regional scale. Thus, rainfall may be a good predictor of actual stocking rates. However, rainfall data are scarce in many regions, and their spatial resolution is usually much coarser than needed to set or to evaluate wildlife or livestock stocking rates. We here show a relationship between livestock biomass and an annual vegetation index (normalized-difference vegetation index-integrated value, NDVI-I) calculated from re- motely sensed data on spectral properties of rangelands of Argentina. The relationship is as strong or even stronger than previously reported correlations between herbivore biomass and rainfall. This, together with the greater availability and higher spatial resolution of satellite data, makes remote sensing a potentially valuable tool to predict stocking rates for regions, landscapes, and different portions of a landscape. The form of the relationship between stocking rate and average NDVI-I was exponential, which, as previously shown, indicates an increasing herbivore load per unit of primary production as rainfall or pro- ductivity increases. This may be at least partially explained by the fact that the NDVI interannual variation and seasonality were negatively related with average NDVI-I. Thus, stocking rate may increase exponentially because of an increasing year-to-year reliability of the forage resource and a more even distribution within the year.

Effect of defoliation intensity on aboveground and belowground relative growth rates

According to a simple growth model, grazed and ungrazed plants may have equal absolute growth rates provided that the relative growth rate (RGR) of grazed plants increases exponentially with grazing intensity (proportion of biomass removed). This paper reports results from an experiment designed to determine whether plants of two grass species subjected to a gradient of defoliation intensities, from 0 to 100% aboveground biomass removal, showed such a response. The relationship between aboveground RGR and defoliation intensity was exponential and closely matched the theoretical relationship of equal absolute growth rate. Thus, plants showed the same aboveground growth regardless of defoliation intensity thanks to an exponential stimulation of RGR by defoliation. Belowground RGR was depressed by defoliation of more than 20% of the above-ground biomass. In spite of the drastic modification imposed by the treatments on the relative proportions of different plant parts, after a 42-day recovery period basic allometric relationships, such as root:shoot and leafarea: weight ratios, were not affected by defoliation intensity. Exponential aboveground compensatory responses represent a key feedback process resulting in constant aboveground growth regardless of defoliation intensity and appear to be a simple consequence of strong commitments to certain allometric relationships.