Rodolfo A. Golluscio

Resource partitioning between shrubs and grasses in the Patagonian steppe

SummaryExperiments were conducted in the Patagonian steppe in southern South America to test the following hypotheses: (a) grasses take up most of the water from the upper layers of the soil and utilize frequent and short-duration pulses of water availability; (b) shrubs, on the contrary, take up most of the water from the lower layers of the soil and utilize infrequent and long-duration pulses of water availability. Grasses and shrubs were removed selectively and the performance of plants and the availability of soil resources were monitored. Results supported the overall hypothesis that grasses and shrubs in the Patagonian steppe use mainly different resources. Removal of shrubs did not alter grass production but removal of grasses resulted in a small increase in shrub production which was mediated by an increase in deep soil water and in shrub leaf water potential. The efficiency of utilization of resources freed by grass removal was approximately 25%. Shrubs used water exclusively from lower soil layers. Grasses took up most of the water from upper layers but they were also capable of absorbing water from deep layers. This pattern of water partitioning along with the lack of response in leaf nitrogen to the removal treatments suggested that shrubs may be at a disadvantage to grasses with respect to nutrient capture and led to questions about the role of nutrient recirculation, leaching, and nitrogen fixation in the steppe.

Differential use of large summer rainfall events by shrubs and grasses: a manipulative experiment in the Patagonian steppe

Abstract In the Patagonian steppe, years with above-average precipitation (wet years) are characterized by the occurrence of large rainfall events. The objective of this paper was to analyze the ability of shrubs and grasses to use these large events. Shrubs absorb water from the lower layers, grasses from the upper layers, intercepting water that would otherwise reach the layers exploited by shrubs. We hypothesized that both life-forms could use the large rainfalls and that the response of shrubs could be more affected by the presence of grasses than vice versa. We performed a field experiment using a factorial combination of water addition and life-form removal, and repeated it during the warm season of three successive years. The response variables were leaf growth, and soil and plant water potential. Grasses always responded to experimental large rainfall events, and their response was greater in dry than in wet years. Shrubs only used large rainfalls in the driest year, when the soil water potential in the deep layers was low. The presence or absence of one life-form did not modify the response of the other. The magnitude of the increase in soil water potential was much higher in dry than in humid years, suggesting an explanation for the differences among years in the magnitude of the response of shrubs and grasses. We propose that the generally reported poor response of deep-rooted shrubs to summer rainfalls could be because (1) the water is insufficient to reach deep soil layers, (2) the plants are in a dormant phenological status, and/or (3) deep soil layers have a high water potential. The two last situations may result in high deep-drainage losses, one of the most likely explanations for the elsewhere-reported low response of aboveground net primary production to precipitation during wet years.

Plant functional types and ecological strategies in Patagonian forbs

Abstract. We identified four major functional types of forbs in the Patagonian steppe, taking into account phenological and morphological traits: (1) shallow-rooted mesophytic species (annuals), (2) shallow-rooted non-mesophytic species, (3) deep-rooted evergreen species, and (4) deep-rooted deciduous species. The major attributes differentiating these groups were the date at which seasonal growth ended, rooting depth, sprouting depth, distance between shoots of the same plant, and degree of ‘mesophytism’. We used Cluster and Principal Components Analyses to identify the groups, and the attributes determining them. Late-growth-cycle types had deep roots and/or high ‘xerophytism’. Late-cycle-xerophytic types had a great sprouting depth, and late-cycle-deep-rooted types had a great distance between shoots of the same plant. On the basis of current knowledge of the structure and functioning of the Patagonian steppe, we suggested three explanations to account for these correlations. 1. Late-cycle forbs survive summer water deficit if they have xerophytic characteristics that reduce transpiration water losses, and/or they have deep roots that increase water uptake. 2. Sprouting depth results from the shift of active buds to dormant buds at the end of the cycle. Summer forbs have a great sprouting depth because only buds which are located deep in the soil survive hot and dry summers. 3. Distant shoots of summer forbs allow them simultaneously to use the high protection against desiccating winds provided by shrubs, and the ample water availability of bare soil patches. All the functional types of forbs depend on winter water recharge to begin their cycles, but each one completes its cycle by using a different portion of the water resources available in spring and summer.

Woody and herbaceous aboveground production of a Patagonian steppe.

Aboveground net primary production (ANPP) of the Patagonian steppe in southwestern Chubut (Argentina) was estimated using a harvest technique to assess the herbaceous (mainly grass) component and a double sampling technique to evaluate shrub production. The latter requires the measurement of plant dimensions and the harvest of shrub biomass in small plots. This technique, by virtue of having an explicit biological model which considers both shrub size and production per unit surface of plant, allows comparisons among years, sites, and treatments. Detailed estimates of ANPP yielded a value of 79 g of dry matter (DM) m-2 yr-1 (SE = 19 g DM m-2 yr-1) for an annual rainfall of 191 mm. Our estimates fits (+/- 17%) predictions of 4 models relating primary production to annual precipitation. Two thirds of production were accounted for by perennial grasses and one third by shrubs. A less detailed method, which uses only peak biomass, gave ANPP estimates for 4 additional years ranging from 21 to 75 g DM m-2 yr-1 while annual precipitation during this period ranged from 55 to 167 mm. There was a large reduction in ANPP during a year of extreme drought; however, there were no increases in ANPP during years with above-average precipitation. This suggests that the carrying capacity for the Patagonian steppe may not be linearly related to precipitation.

Soil water availability in the Patagonian arid steppe: Gravel content effect

Abstract A model for estimating volumetric water content as a function of soil water potential and gravel content was developed. In the system studied gravel can withhold up to 67% of the amount held by the fine material. Water content at field capacity decreases 50% when gravel content (>11 mm) increases from 0% to 40% of total soil weight. For those values of gravel content, penetration depth of a rainfall of 20 mm increases from 38 cm to 59 cm when the initial water content is 60% of the field capacity.

Sheep Grazing Decreases Organic Carbon and Nitrogen Pools in the Patagonian Steppe: Combination of Direct and Indirect Effects

We explored the net effects of grazing on soil C and N pools in a Patagonian shrub–grass steppe (temperate South America). Net effects result from the combination of direct impacts of grazing on biogeochemical characteristics of microsites with indirect effects on relative cover of vegetated and unvegetated microsites. Within five independent areas, we sampled surface soils in sites subjected to three grazing intensities: (1) ungrazed sites inside grazing exclosures, (2) moderately grazed sites adjacent to them, and (3) intensely grazed sites within the same paddock. Grazing significantly reduced soil C and N pools, although this pattern was clearest in intensely grazed sites. This net effect was due to the combination of a direct reduction of soil N content in bare soil patches, and indirect effects mediated by the increase of the cover of bare soil microsites, with lower C and N content than either grass or shrub microsites. This increase in bare soil cover was accompanied by a reduction in cover of preferred grass species and standing dead material. Finally, stable isotope signatures varied significantly among grazed and ungrazed sites, with δ15N and δ13C significantly depleted in intensely grazed sites, suggesting reduced mineralization with increased grazing intensity. In the Patagonian steppe, grazing appears to exert a negative effect on soil C and N cycles; sound management practices must incorporate the importance of species shifts within life form, and the critical role of standing dead material in maintaining soil C and N stocks and biogeochemical processes.

Range assessment using remote sensing in Northwest Patagonia (Argentina).

A methodology based on remotely sensed data (LANDSAT MSS) was used for rapid assessment of rangelands in the grass and shrub steppes of NW Patagonia (Argentina). We calibrated Normalized Difference Vegetation Index (NDVI) data using total plant cover, grass cover, shrub cover, and floristic data. Total vegetation cover and grass cover was predicted with high accuracy from Normalized Difference Vegetation Index data. The correlation between observed and estimated cover was 0.87 and 0.82 (P < 0.01) for total cover and grass cover respectively. The correlation was lower for shrub cover than for grass (r = 0.45, p < 0 .01). Normalized Difference Vegetation Index data was used to accurately predict cover of Festuca pallescens (St. Yves) Parodi (coiron blanco) and Nassau via glomerulosa (Lag.) Don (colapiche), 2 species with contrasting response to grazing in the Occidental district of Patagonia, and typical of vegetation with very different grazing values. The correlation between observed and estimated from the Normalized Difference Vegetation Index cover was 0.67 and 0.53 (P < 0.01) for Festuca pallescens (coiron blanco) and Nassauvia glomerulosa (colapiche) respectively.

Aridity and grazing as convergent selective forces: an experiment with an Arid Chaco bunchgrass

It has been proposed that aridity and grazing are convergent selective forces: each one selects for traits conferring resistance to both. However, this conceptual model has not yet been experimentally validated. The aim of this work was to experimentally evaluate the effect of aridity and grazing, as selective forces, on drought and grazing resistance of populations of Trichloris crinita, a native perennial forage grass of the Argentinean Arid Chaco region. We collected seeds in sites with four different combinations of aridity and grazing history (semiarid/ subhumid x heavily grazed/lightly grazed), established them in pots in a common garden, and subjected the resulting plants to different combinations of drought and defoliation. Our results agreed with the convergence model. Aridity has selected T. crinita genotypes that respond better to drought and defoliation in terms of sexual reproduction and leaf growth, and that can evade grazing due to a lower shoot: root ratio and a higher resource allocation to reserves (starch) in stem bases. Similarly, grazing has selected genotypes that respond better to drought and defoliation in terms of sexual reproduction and that can evade grazing due to a lower digestibility of leaf blades. These results allow us to extend concepts of previous models in plant adaptation to herbivory to models on plant adaptation to drought. The only variable in which we obtained a result opposite to predictions was plant height, as plants from semiarid sites were taller (and with more erect tillers) than plants from subhumid sites; we hypothesize that this result might have been a consequence of the selection exerted by the high solar radiation and soil temperatures of semiarid sites. In addition, our work allows for the prediction of the effects of dry or wet growing seasons on the performance of T. crinita plants. Our results suggest that we can rely on dry environments for selecting grazing-resistant genotypes and on high grazing pressure history environments for selecting drought-resistant ones.

Environmental controls of NDVI dynamics in Patagonia based on NOAA-AVHRR satellite data

. Variation in vegetation in extra-Andean Patagonia (Argentina) was analyzed using spectral data derived from AVHRR/NOAA satellite. The study of seasonal dynamics of the Normalized Difference Vegetation Index (NDVI, i.e. a combined index of the reflection in the red and infrared bands) highlighted similarities in functional aspects between regional vegetation units which are dissimilar in a geographical, physiognomical and/or floristical way, and also suggested that gross primary production is correlated with mean annual rainfall. The first axis in a Principal Component Analysis of NDVI data was correlated (r2 = 0.90) with NDVI as integrated for the study period. The second axis was correlated (r2 = 0.50) with the differences in NDVI during the growing season, reflecting seasonality. Mean annual rainfall accounted for 60% of integrated NDVI variability among vegetation units. Much of the residual variance (62%) was accounted for by the inverse of the distance to the Atlantic Ocean, which is interpreted as an ocean effect on vegetation functioning in the extra-Andean Patagonia.

The Use of Satellite Imagery in Quantitative Phytogeography: A Case Study of Patagonia (Argentina)

Ecological surveys of arid zones, both with basic and applied phytogeographical objectives, are very difficult because of their magnitude and the low density of their renewable natural resources. The lack of adequate knowledge of the heterogeneity and the functioning of the vegetation is common in these regions. This leads to increasing damage of the environment as a consequence of the inbalance between supply and demand in the use of their natural resources (Soriano 1986).