Mónica B. Bertiller

Soil nitrogen dynamics in northeastern Patagonia steppe under different precipitation regimes

Small-scale heterogeneity of plant cover and highly variable precipitation events in dry regions can strongly influence N dynamics. We evaluated the differences in N availability (Ni), N mineralization (Nmin), flush of microbial-N (N-MF) and soil moisture (SM) at 0–20 cm depth among four types of patches characteristic of heavily grazed areas in the northeastern Patagonia steppe of Larrea divaricata and Stipa spp. Soil samples were taken monthly during two years of differing annual precipitation (178 mm in 1994 and 325 mm in 1995). Ni and SM were also measured at 20–40 cm depth. Additionally, we estimated the potential N mineralization (pNmin) during two months in both winter and summer in laboratory incubations at 20% soil moisture and 25°C. Sampled patches included: undisturbed patches of shrubs and perennial grasses (GSP), incipient patches of Larrea divaricata and perennial grasses (IGSP), incipient patches of the perennial grass Stipa tenuis (GP), and bare soil (BS). Mineralization rates were much higher during the wet year, and higher in GSP and IGSP than in GP and BS. The prevailing form of Ni was NH4+–N, but pulses of NO3-–N were measured in field incubations when SM was higher than 10%; NO3-–N was also the main form of Ni in pNmin assays. Flush of microbial-N depended mainly on plant cover, following the sequence: GSP>IGSP>GP=BS. It was not correlated with soil moisture, except in the GSP patches, and exhibited lower values during the wet year. Available N (as NH4+–N) was higher in the subsurface than in the surface samples during the wet year. The relative importance of N-MF and Nmin as indicators of spatial and temporal changes in N dynamics, and the role of deep-rooted shrubs in the recovery of soil N fertility, are discussed.

Grazing effects on sustainable semiarid rangeiands in Patagonia: The state and dynamics of the soil seed bank

The composition of the germinable seed bank was studied in four vegetation states of the Festuca pallescens grasslands in semiarid Patagonia during four years. The aim of this study was to test whether aboveground vegetation states resulting from grazing exclusion or different combinations of grazing and topography are reflected in different states of the germinable seed bank. The size of the total and dicot germinable seed bank was positively related to the total cover in each state. Dicots dominated all germinable seed bank states. Carex patagonica increased its cover as well as its germinable seed bank under grazing disturbance. Grazing did not reduce the germinable seed bank of perennial grasses in uplands where the grazing pressure is lower as compared with slopes. In slopes the germinable seed bank of perennial grasses was significantly reduced by grazing. A reduction of the length of the grazing period in late spring increases the germinable seed bank of perennial grasses both in upland and slope. These results are interpreted in the frame of a model of management techniques where grazing exclusion during late spring and late summer increases the seed bank of the perennial grasses and promotes their establishment in uplands. The artificial addition of seeds of perennial grasses and the manipulation of the soil surface in order to increase “safe sites” appear as management alternatives that deserve further evaluation to improve plant reestablishment in slopes.

Plant phenology, leaf traits and leaf litterfall of contrasting life forms in the arid Patagonian Monte, Argentina

Abstract Question: Do coexisting plant life forms differ in overall phenology, leaf traits and patterns of leaf litterfall? Location: Patagonian Monte, Chubut Province, Argentina. Methods: We assessed phenology, traits of green and senesced leaves and the pattern of leaf litterfall in 12 species of coexisting life forms (perennial grasses, deciduous shrubs, evergreen shrubs). Results: We did not identify differences in phenology, leaf traits and patterns of leaf litterfall among life forms but these attributes contrasted among species. Independent of the life form, the maintenance of green leaves or vegetative growth during the dry season was mostly associated with leaves with high leaf mass per area (LMA) and high concentration of secondary compounds. Low LMA species produced low litterfall mass with low concentration of secondary compounds, and high N concentration. High LMA species produced the largest mass of leaf litterfall. Accordingly, species were distributed along two main dimensions of ecological variation, the dimension secondary compounds in leaves - length and timing of the vegetative growth period (SC – VGP) and the dimension leaf mass per area - leaf litterfall mass (LMA – LLM). Conclusions: Phenology, leaf traits and leaf litterfall varied among species and overlapped among life forms. The two dimensions of ecological variation among species (SC – VGP, LMA – LLM) represent distinct combinations of plant traits or strategies related to resource acquisition and drought tolerance which are reflected in the patterns of leaf litterfall. Nomenclature: Correa (1971, 1978, 1984, 1988, 1999).

Relationship between plant nitrogen conservation strategies and the dynamics of soil nitrogen in the arid Patagonian Monte, Argentina

During three consecutive years with contrasting precipitation, we analysed the relationship between strategies of N conservation in the dominant plant functional groups (perennial grasses and evergreen shrubs) of the Patagonian Monte and the main components of N cycling in soil. We hypothesised that the different patterns of N conservation in perennial grasses and evergreen shrubs would have direct consequences for soil-N, inorganic-N release and microbial-N flush in soil. In autumn and late spring of 1999, 2000, and 2001, we assessed N and C concentration in green and senesced leaves, N-resorption efficiency and C/N ratio in senesced leaves of three dominant species of each plant functional group. In the soil associated with species of each plant functional group, we determined N and C concentration, potential-N mineralisation, and the associated microbial-N flush. Slow-growing evergreen shrubs exhibited low N-concentration in green leaves, high N-concentration in senesced leaves and low N-resorption from senescing leaves. In contrast, fast-growing perennial grasses showed high N-concentration in green leaves, low N-concentration in senesced leaves, and high N-resorption from senescing leaves. In evergreen shrubs, the maintenance of long-lasting green leaves with low N-concentration was the most important mechanism of N conservation. In contrast, perennial grasses conserved N through high N-resorption from senescing leaves. Soil-N concentration, potential N-mineralisation, and microbial-N flush in the soil were higher underneath evergreen shrubs than beneath perennial grasses. Observed differences, however, were lower than expected considering the quality of the organic matter supplied by each plant fuctional group to the soil. A possible reason for this relatively weak trend may be the capacity of evergreen shrubs to slow down N cycling through low leaf turnover and the presence of secondary compounds in leaves. Alternatively or simultaneously, the weak relationship between plant and soil N could result from shrubs being able to colonise N-poor soils while grasses may preferably occupy fertile microsites previously influenced by the decomposition pathway of evergreen shrubs. Differences between evergreen shrubs and perennial grasses in the mechanisms of plant N-conservation and in components of N cycling in the underlying soil were consistent over the three years of the study with differing precipitation. Inter-annual differences in N concentration in green leaves and in the microbial-N flush in soil indicate that during the wettest year fast-growing perennial grasses would outcompete slow-growing evergreen shrubs and microorganisms for N uptake.

Seasonal variation in the seed bank of a Patagonian grassland in relation to grazing and topography

. The germinable seed bank of Festuca spp., Carex patagonica, and other annuals and perennials in a semiarid Patagonian grassland was analyzed every three months. The effects of grazing, topography and microsites with respect to established grass tussocks on the germinable-seed bank were also analyzed. The total germinable-seed bank was larger in summer after seed rain. At this time of the year most of the seeds were from annuals and perennial dicots. Seeds of the dominant Festuca spp. were the main components of the graminoid seed bank, which was homogeneously distributed in patches of bare soil. In spring, i.e. some months after the seed rain, the germinable-seed bank of most of the species was greatly reduced, while the seed bank of Carex patagonica did not change significantly. At this time of the year, the graminoid seed bank was heterogeneously distributed in space, with the seeds accumulating in wind-protected microsites. In the grazing treatments, the seed bank of the perennial grasses was reduced proportionally to the decrease of the plant cover. On the other hand, the germinable-seed bank of C. patagonica increased with the grazing treatments, in relation to the increase of vegetation cover. Topographical position had two types of effect on the seed bank of perennial grasses. One was the increase of the germinable-seed bank, after the seed rain, on the relatively warmer northeast facing slopes, which was related to a higher seed rain in these places. The other was an increased seed loss on slopes. The different seed-bank strategies observed in this grassland are compared and discussed in relation to strategies described for other grasslands.

Patterns of nitrogen conservation in shrubs and grasses in the Patagonian Monte, Argentina.

We analysed the main plant strategies to conserve nitrogen in the Patagonian Monte. We hypothesized that the two main plant functional groups (xerophytic evergreen shrubs and mesophytic perennial grasses) display different mechanisms of nitrogen conservation related to their structural and functional characteristics. Evergreen shrubs are deep-rooted species, which develop vegetative and reproductive growth from spring to late summer coupled with high temperatures, independently from water inputs. In contrast, perennial grasses are shallow-rooted species with high leaf turnover, which display vegetative growth from autumn to spring and reproductive activity from mid-spring to early-summer, coupled with precipitation inputs. We selected three evergreen shrubs (Larrea divaricata Cav., Atriplex lampa Gill. ex Moq. and Junellia seriphioides (Gilles and Hook.) Moldenke) and three perennial grasses (Stipa tenuis Phil., S. speciosa Trin. and Rupr. and Poa ligularis Nees ex Steud.), characteristic of undisturbed and disturbed areas of the Patagonian Monte. N concentration in expanded green and senesced leaves was estimated in December 1997 (late spring) and June 1998 (late autumn). Deep-rooted evergreen shrubs displayed small differences in N concentration between green and senesced leaves (low N-resorption efficiency), having high N concentration in senesced leaves (low N-resorption proficiency). Shallow-rooted perennial grasses, conversely, showed high N-resorption efficiency and high N-resorption proficiency (large differences in N concentration between green and senesced leaves and very low N concentration in senesced leaves, respectively). The lack of a strong mechanism of N resorption in evergreen shrubs apparently does not agree with their ability to colonize N-poor soils. These results, however, may be explained by lower N requirements in evergreen shrubs resulting from lower growth rates, lower N concentrations in green leaves, and lower leaf turnover as compared with perennial grasses. Long-lasting N-poor green tissues may, therefore, be considered an efficient mechanism to conserve N in evergreen shrubs in contrast with the mechanism of strong N resorption from transient N-rich tissues displayed by perennial grasses. Evergreen shrubs with low N-resorption efficiency provide a more N-rich substrate, with probably higher capability of N mineralization than that of perennial grasses, which may eventually enhance N fertility and N availability in N-poor soils.

Leaf strategies and soil N across a regional humidity gradient in Patagonia

We analyzed leaf traits related to carbon-fixation, nutrient conservation strategies, and decomposability and their relationships with potential N-mineralization and microbial N in soil in 19 species of 5 dominant life forms growing in 40 sites across a regional humidity gradient in northern Patagonia. We hypothesized that (1) the shifting of species and life forms across the humidity gradient is related to a shifting in traits of green and senesced leaves with some overlapping among life forms, and (2) leaf traits associated with litter decomposition are related to the potential dynamics of soil-N across the humidity gradient. LMA in green leaves and P-resorption efficiency decreased with humidity while concentrations of lignin and total phenolics in green and senesced leaves and P concentration in senesced leaves increased with humidity. Soil C and N concentrations were positively correlated to humidity. Increasing soil N concentration was related to increasing rates of absolute (per unit soil mass) potential net N-mineralization and microbial-N flush. Relative (per unit N mass) potential net N-mineralization and microbial-N flush decreased with soil N and were inversely correlated to lignin concentration and C/N ratio in senesced leaves. We found overlapping in N concentration and C/N ratio in green and senesced leaves, P concentration in green leaves, and N resorption among species and life forms across the humidity gradient. We concluded that (1) leaf traits related to carbon fixation and the decomposition pathway significantly varied with humidity and were not overlapped between plant life forms from dry and humid habitats, (2) the largest overlapping among species and plant life forms across the gradient occurred in those leaf traits related to N conservation in the plant, and (3) life forms from humid habitats produce more recalcitrant litter that induce lower rates of relative potential net N mineralization (per unit N) than those of dry habitats.

Spatial sex segregation in the dioecious grass Poa ligularis in northern Patagonia: the role of environmental patchiness

We examined the effect of environmental patchiness on the spatial segregation of the sexes in the dioecious anemophilus grass Poa ligularis. Because the species is sensitive to grazing, a better understanding of environmental factors that control its spatial distribution and abundance could improve conservation efforts. We hypothesized that (i) males and females are spatially segregated in the microenvironments created by plant patches as the result of sexual specialization in habitat and/or resources use, (ii) sexual specialization is related to different tolerance to competition and reproductive costs of males and females, and (iii) changes in patch structure affect the microenvironment and the intensity of spatial segregation of the sexes. We analyzed the spatial distribution of sexes at three sites with different plant and micro-environmental patchiness and performed a controlled competition experiment with different substitution of males and females. Our results showed that large plant patches created larger sheltered soil fertility islands than small patches. As patch size and their area of influence increased, the density and the spatial segregation of the sexes of P. ligularis also increased, resulting in biased habitat-specific sex ratios. In accordance with their higher reproductive costs, females were more frequent in sheltered (low air evaporative demand) and nitrogen-rich areas inside patch perimeters than males. Females were also better able to tolerate inter-sexual competition than males. In contrast, males tolerated low nitrogen concentration in soil and low sheltering, probably gaining advantage in pollen dispersal. Inter- and intra-sexual competition, however, affected the reproductive output of both sexes. From the point of view of conservation, environmental patchiness is important to the status of P. ligularis populations. The reduction of patch size limits the available microsites, biases the sex ratio towards males inside patches, increases inter- and intra-sexual competition, and it might be expected to decrease overall seed and pollen production and consequently potential recruitment.

Seedling emergence and survival in contrasting soil microsites in Patagonian Monte shrubland

The potential of two perennial species (Larrea divaricata and Stipa tenuis) to colonize different soil microsites was analyzed in the Patagonian Monte shrubland. We hypo- thesize that the short-lived grass S. tenuis is more able to colonize the soil of microsites beneath vegetation patches where N-fertility is higher than those in bare soil, while the long-lived shrub L. divaricata colonizes different soil microsites irrespective of their N fertility. A greenhouse experiment was carried out to evaluate the emergence and survival of both species in different soil microsites at different water, inorganic N and seed densities. In all cases soil microsites were seed limited since the addition of viable seeds increased seedling emergence. Both species showed, however, different abilities to emerge and survive in different soil microsites. Microsites of bare soil were more favourable for seedling emergence and survival ofL. divaricata than those beneath vegetation patches, independent of their water status. This ability of L. divaricata can not be explained on the basis of increased water or N availability, but probably because of lower salt content of bare soil. The addition of inorganic N reduced the survival of L. divaricata in both microsites but increased individual plant performance. The emergence and survival of S. tenuis was not different in both types of soil microsites but the addition of inorganic N increased seedling emergence and plant biomass. According to these results, emergence and plant performance of S. tenuis may be promoted during humid years by increased concentration of inorganic N. Since N mineralization occurs at a higher rate in soil microsites beneath vegetation patches than in those of bare soil, higher plant performance and probably establishment of S. tenuis is to be expected. These results are consistent with an existing conceptual model of plant dynam- ics under various grazing intensities in the Patagonian Monte shrubland based on previous field observations.

Environmental patterns and plant distribution along aprecipitation gradient in western Patagonia

Abstract Plant species distribution along a gradient of precipitation was analysed inwestern Patagonia. First, the affinity of 53 vegetation releves was tested by principal component analysis of the matrix of n × n binary correlations among species cover values. Further, the relationship between plant cover (Festuca pallescens, shrubs, and Stipa spp.) and the environmental factors (precipitation, soil texture, soil depth, altitude, landscape form, aspect, and disturbance) was analysed by stepwise regression analysis. Five homogeneous vegetation units could be identified at several different regions of the gradient. Soil characteristics associated to precipitation levels (i.e. texture, profile depth), but not precipitation per se, were strongly related to the cover of F. pallescens and shrub species. A model of transitions in the composition of vegetation is introduced which includes soil variables, site conditions and the spatial distribution of grazing herds.