Forage quality in relation to long-term grazing history,current-year defoliation,and water resource

Forage nitrogen concentrations, nitrogen yields, and in vitro digestibilities were assessed in shortgrass steppe that had been ungrazed, lightly, or heavily grazed for 50 years. Caged plots were defoliated in amounts based upon removals observed in naturallygrazed reference plots or not defoliated. This was done in a year of average precipitation and with a supplemental water treatment to simulate a wet year. In general, current-year defoliation had positive effects, and longterm grazing and supplemental water had negative effects, on forage nitrogen concentrations and digestibilities. However, defoliation interacted with long-term grazing in determning forage nitrogen concentrations, and with grazing and with watering in determining digestibilities. Nitrogen concentration and digestibility increased with defoliation in lightly, but not in heavily, grazed treatments. The dilution effect of supplemental water an digestibilities through increased plant growth was offset by defoliation. The negative effects of long-term grazing on forage quality were small, equally or more than compensated for by defoliation in a year of average precipitation, but more pronounced in the simulated wet year. Nitrogen yields and digestible forage production were usually increased by defoliation, but this depended upon grazing and watering treatments. Increased nitrogen and digestible forage yields and concentrations in response to defoliation were greater than the biomass response in lightly grazed grassland. For both nitrogen and digestibility, yields were greater in grazed than ungrazed treatments in the year of average precipitation, but less in the simulated wet year. Optimizing quantity and year-to-year stability of nitrogen and digestible forage yield may best be achieved with light grazing rather than no or heavy grazing. Clipping was conducted in a manner closely resembling the natural pattern and intensity of defoliation by the cattle, and confirms the potential for a positive feedback of increased forage quality with defoliation observed in pot experiments. Long-term heavy grazing can diminish this response. Quantily (aboveground primary production, ANPP), quantity of quality (digestible and N yields), and quality (concentrations) do not necessarily respond similarly in interactions between current-year defoliation, long-term grazing history, and level of water resource.     

Ecological conditions that determine when grazing stimulates grass production

Summary We report the results of a pot experiment that examined the effects of three ecologically important factors controlling plant growth rates in savanna grasslands: defoliation, soil nitrogen and soil water availability. The experiment was conducted in the Amboseli region in east Africa, and was designed to simulate natural conditions as far as possible, using local soils and a grass species that is heavily grazed by abundant large herbivores. Productivity by different plant components was reduced, stimulated or unchanged by defoliation, depending on specific watering and fertilization treatments. Total above-ground production was stimulated by defoliation and was maximized at moderate clipping intensities, but this was statistically significant only when plants were watered infrequently (every 8 days), and most important, periods between clipping events were extended (at least 24 days). Under these conditions, plant growth rates were limited by water availability at the time of clipping, and soil water conserved in clipped, compared to unclipped plants. Within a given fertilization treatment, whole-plant production was never stimulated by defoliation because root growth was unaffected or inhibited by clipping. However, when fertilization was coupled to defoliation, as they are in the field, whole-plant production by fertilized and moderately clipped plants exceeded production by infertilized, unclipped plants. Under this interpretation, maximum whole-plant production coincided with optimum conditions for herbivores (maximum nitrogen concentration in grass leaves) when watering was frequent, and plants were moderately defoliated. However, these conditions were not the same as those that maximized relative above-ground stimulation of growth (infrequent watering and clipping).The results indicate that above-ground grass production can be stimulated by grazing, and when that is likely to occur. However, the results emphasize that plant production responses to defoliation can vary widely, contigent upon a complex interaction of ecological factors.     

Growth responses of arctic graminoids following grazing by captive lesser snow geese

The effects of grazing by captive goslings of the Lesser Snow Goose on coastal vegetation at La Pérouse By. Manitoba were investigated. Swards of Carex subspathacea, Festuca rubra and Calamagrostis deschampsioides were grazed once for different periods (0–180 min) and regrowth of vegetation determined, based on measurements of standing crop, net above-ground primary production (NAPP) and forage quality (leaf nitrogen content). The amounts of foliage removed from swards of Carex subspathacea increased with the length of the grazing period, but after 44 days of regrowth there were no significant differences in above-ground biomass between control and grazed plots. While the amount of foliage removed by goslings from swards of Festuca rubra increased with the length of the grazing period (except after 150 min of grazing), the increase in biomass following defoliation was similar among treatments. Goslings removed little biomass from swards of Calamagrostis deschampsioides, even when the opportunity for grazing was 180 min. No significant differences in standing-crop or NAPP between grazed and ungrazed plots were detected by the end of summer. Grazing had no significant effect on amounts of nitrogen in leaf tissue of all species, suggesting that faecal nitrogen was not rapidly incorporated into plant biomass within the growing season. Patterns of regrowth of these species are compared to that of Puccinellia phryganodes. An increase in goose numbers in recent years has led to birds foraging on less preferred species, such as Calamagrostis deschampsiodes and Festuca rubra. Their poor nutritional quality and a lack of a rapid growth response following defoliation may explain, in part, the decline in the weight of wild goslings recorded over the last decade.     

Effect of grazing on community structure and productivity of a Uruguayan grassland

Grasslands and their grazers provide some of the most compelling examples for studying the relationship between diversity, productivity, and disturbance. In this study, we analyzed the impact of grazing-induced changes in species composition and community structure upon the productivity of a grassland in the Campos region, Uruguay. We compared three treatments: a continuously grazed area, a 9-year old exclosure to domestic herbivores, and grazing-simulated plots inside the exclosure, which were clipped so that their standing biomass resembled that of the grazed area. We studied the community composition of the grazed and ungrazed situations, and determined biomass and above-ground net primary production (ANPP) of the three treatments during 1 year. Grazed plots had higher species richness and diversity than the exclosure. Grazing resulted in the replacement of some cool-season, tussock grasses by warm-season, prostrate grasses. ANPP was 51% higher under grazing than in the exclosure, but the grazing-simulated plots inside the exclosure were the most productive treatment, 29% higher than the grazed plots. Thus, two components of grazing effect may be postulated for this grassland. The structural component resulted in higher ANPP, probably due to the elimination of standing dead biomass. The species composition component resulted in lower ANPP once the structural component was controlled, probably due to the shift to warm-season phenology and prostrate habit. Our findings contrast with a similar experiment carried out in the neighbouring Flooding Pampa region, which suggests that the relationship between grazing and community structure and function is difficult to generalize.     

Responses of soil respiration to simulated precipitation pulses in semiarid steppe under different grazing regimes

Aims Precipitation pulses and different land use practices (such as grazing) play important roles in regulating soil respiration and carbon balance of semiarid steppe ecosystems in Inner Mongolia. However, the interactive effects of grazing and rain event magnitude on soil respiration of steppe ecosystems are still unknown. We conducted a manipulative experiment with simulated precipitation pulses in Inner Mongolia steppe to study the possible responses of soil respiration to different precipitation pulse sizes and to examine how grazing may affect the responses of soil respiration to precipitation pulses.Methods Six water treatments with different precipitation pulse sizes (0, 5, 10, 25, 50 and 100 mm) were conducted in the ungrazed and grazed sites, respectively. Variation patterns of soil respiration of each treatment were determined continuously after the water addition treatments.Important findings Rapid and substantial increases in soil respiration occurred 1 day after the water treatments in both sites, and the magnitude and duration of the increase in soil respiration depended on pulse size. Significantly positive relationships between the soil respiration and soil moisture in both sites suggested that soil moisture was the most important factor responsible for soil respiration rate during rain pulse events. The ungrazed site maintained significantly higher soil moisture for a longer time, which was the reason that the soil respiration in the ungrazed site was maintained relatively higher rate and longer period than that in the grazed site after a rain event. The significant exponential relationship between soil temperature and soil respiration was found only in the plots with the high water addition treatments (50 and 100 mm). Lower capacity of soil water holding and lower temperature sensitivity of soil respiration in the grazed site indicated that degraded steppe due to grazing might release less CO₂ to the atmosphere through soil respiration under future precipitation and temperature scenarios.     

Effects of bison grazing on Andropogon gerardii and Panicum virgatum in burned and unburned tallgrass paririe

Summary Responses to clipping and bison grazing in different environmental contexts were examined in two perennial grass species, Andropogon gerardii and Panicum virgatum, on the Konza Prairie in northeastern Kansas. Grazed tillers had lower relative growth rates (RGR) than clipped tillers following defoliation but this difference was transient and final biomass was not affected by mode of defoliation. Grazed tillers of both species had higher RGR throughout the season than ungrazed tillers, resulting in exact compensation for tissue lost to defoliation. However, A. gerardii tillers which had been grazed repeatedly the previous year (1988) had reduced relative growth rates, tiller biomass and tiller survival in 1989. This suggests that the short-term increase in aboveground relative growth rates after defoliation had a cost to future plant growth and tiller survival.In general, the two species had similar responses to defoliation but their responses were altered differentially by fire. The increase in RGR following defoliation of A. gerardii was relatively greater on unburned than burned prairie, and was influenced by topographic position. P. virgatum responses to defoliation were similar in burned and unburned prairie. Thus grazing, fire, and topographical position all interact to influence tiller growth dynamics and these two species respond differently to the fire and grazing interaction. In addition, fire may interact with grazing pattern to influence a plants' grazing history and thus its long-term performance.     

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.     

Interactive regulation of grass yield and chemical properties by defoliation,a salivary chemical,and inorganic nutrition

Summary Clones of 2 C₄ grass species, Sprobolus ioclados and S. pyramidalis, were obtained from more and less heavily grazed grasslands, respectively, in Tanzania's Serengeti National Park. Plants were grown in a factorial experiment to determine the effects of severe defoliation, nutrient limitation, and a salivary chemical (thiamine) on plant growth, nitrogen content, and non-structural carbohydrate content. The experimental design included: (1) species; (2) clipping, with plants either unclipped or clipped weekly to a height of 5 cm; (3) 0.2 ml of distilled water of 0.2 ml of 10 ppb aqueous thiamine sprayed on plants from an atomizer after clipping and identical treatments at the same time to unclipped plants; (4) phosphorus (P) at 0.2 or 1 mM; (5) nitrogen (N) at 3 or 15 mM. Clipping was the major variable affecting plant growth. Total and litter yields were reduced to half and residual plant yield was reduced to 30% of the values for unclipped plants. Clipping interacted strongly with other variables since they commonly had minor effects on clipped plants and major effects on unclipped plants. Exceptions to this generalization were generally due to better performance by S. ioclados under clipping. Compared to lower treatment levels, higher treatment levels promoted total yield of unclipped plants by 52% for N, 43% for thiamine, and 33% for P. In general, thiamine had greater effects than P but lesser effects than N. Thiamine promoted yield and modified the chemical balance of plants by promoting carbohydrate (CHO) concentrations and reducing N concentrations. N and P deficiencies promoted CHO accumulation. Clipping promoted the N of leaves and crowns and reduced the N levels in roots. Leaf blade water and N contents were positively correlated with very little scatter. The slope of the line was different for S. ioclados and S. pyramidalis. Leaf blade water and CHO contents were negatively related but there was more scatter and the species could not be distinguished. The species from more heavily grazed grasslands was conspicuously more sensitive to thiamine application. The results indicated that leaf treatment with thiamine, the only likely source of which in natural grasslands is saliva deposited by feeding herbivores, can have major effects on plant yield and metabolic balances at very low application levels. But under defoliation levels as severe as those imposed in this experiment, which reduced above ground plant biomass to a fourth of the level produced by unclipped plants, growth was so strongly limited by defoliation that neither thiamine nor inorganic nutrients affected plant yield residual from clipping. Therefore, whether chemicals such as thiamine that may be introduced onto grass foliage by grazing ungulates and other herbivores will influence the growth of grazed plants will depend upon the grazing intensity associated with the transfer.     

内蒙古温带草原不同放牧强度和围栏封育对凋落物分解的影响

放牧和围封通过影响植物群落结构和土壤微环境来调控草地生态系统的碳循环。该研究在内蒙古温带草原设置轻度放牧后围封、轻度放牧、重度放牧后围封、重度放牧4种样地, 通过测定干旱年(2011年)和湿润年(2012年)地上、地下凋落物产量、质量及其分解速率和土壤养分含量, 分析不同放牧强度对凋落物形成和分解的影响, 以及围栏封育对生态系统恢复的作用。结果表明: 重度放牧地上凋落物产量和分解速率均高于轻度放牧。干旱年轻度放牧样地地下凋落物产量和分解速率高于重度放牧, 湿润年相反。短期围封显著提高了凋落物产量, 轻度放牧样地围封后地上凋落物分解速率和养分循环加快, 而重度放牧样地围封后地上凋落物分解减慢。因此, 与重度放牧相比, 轻度放牧草地的恢复更适合采用围栏封育措施; 而重度放牧草地的恢复可能还需辅以必要的人工措施。降水显著促进地上、地下凋落物形成和分解。地下凋落物的生产和分解受降水年际波动影响较大, 重度放牧草地对降水变化的敏感度比轻度放牧草地高。地上凋落物分解速率与凋落物N含量显著正相关, 与土壤全N显著负相关, 与地上凋落物C:N和木质素:N相关性不大; 地下凋落物分解速率与凋落物C、C:N和纤维素含量显著负相关。该研究结果将为不同放牧强度的草地生态系统恢复和碳循环研究提供理论依据。     

Effects of grazing intensity and grazing exclusion on litter decomposition in the temperate steppe of Nei Mongol,China

Aims Grazing intensity and grazing exclusion affect ecosystem carbon cycling by changing the plant community and soil micro-environment in grassland ecosystems. The aims of this study were: 1) to determine the effects of grazing intensity and grazing exclusion on litter decomposition in the temperate grasslands of Nei Mongol; 2) to compare the difference between above-ground and below-ground litter decomposition; 3) to identify the effects of precipitation on litter production and decomposition. Methods We measured litter production, quality, decomposition rates and soil nutrient contents during the growing season in 2011 and 2012 in four plots, i.e. light grazing, heavy grazing, light grazing exclusion and heavy grazing exclusion. Quadrate surveys and litter bags were used to measure litter production and decomposition rates. All data were analyzed with ANOVA and Pearson’s correlation procedures in SPSS. Important findings Litter production and decomposition rates differed greatly among four plots. During the two years of our study, above-ground litter production and decomposition in heavy-grazing plots were faster than those in light-grazing plots. In the dry year, below-ground litter production and decomposition in light-grazing plots were faster than those in heavy-grazing plots, which is opposite to the findings in the wet year. Short-term grazing exclusion could promote litter production, and the exclusion of light-grazing could increase litter decomposition and nutrient cycling. In contrast, heavy-grazing exclusion decreased litter decomposition. Thus, grazing exclusion is beneficial to the restoration of the light-grazing grasslands, and more human management measures are needed during the restoration of heavy-grazing grasslands. Precipitation increased litter production and decomposition, and below-ground litter was more vulnerable to the inter-annual change of precipitation than above-ground litter. Compared to the light-grazing grasslands, heavy-grazing grasslands had higher sensitivity to precipitation. The above-ground litter decomposition was strongly positively correlated with the litter N content (R² = 0.489, p < 0.01) and strongly negatively correlated with the soil total N content (R² = 0.450, p < 0.01), but it was not significantly correlated with C:N and lignin:N. Below-ground litter decomposition was negatively correlated with the litter C (R² = 0.263, p < 0.01), C:N (R² = 0.349, p < 0.01) and cellulose content (R² = 0.460, p < 0.01). Our results will provide a theoretical basis for ecosystem restoration and the research of carbon cycling.     

Responses to precipitation treatment for <Emphasis Type="Italic">Haloxylon ammodendron</Emphasis> growing on contrasting textured soils

The responses to precipitation of Haloxylon ammodendron (C.A. Mey.) Bunge (Chenopodiaceae), a small xerophilous tree growing on contrasting textured soils, were evaluated under no, natural, and double precipitation treatments during the entire growing season of 2006. The contrasting textured soils are sandy and heavy textured, and both are the original habitat of H. ammodendron at the south edge of Gubantonggute Desert, Central Asia. Photosynthesis, leaf water potential, transpiration, water use efficiency and leaf biomass production were monitored throughout the growing season. Root distribution of H. ammodendron was evaluated at the end of the experiment. Overall, this small tree did not show significant response to a large summer precipitation pulse or precipitation treatments, in terms of photosynthetic carbon assimilation on either soil. The leaf water potential, transpiration, and water use efficiency appeared to be highly sensitive to a large precipitation pulse and precipitation treatments in sandy soil; and leaf biomass production was also much higher for plants in sandy than that of heavy-textured soil. In sandy soil, defoliation occurred when pre-dawn leaf water potential dropped below −3.0 MPa, while in heavy-textured soil, defoliation occurred when pre-dawn leaf water potential dropped below −3.75 MPa. For similar above-ground parts, the small trees at the sandy site developed much deeper root systems and had nearly double the surface area of feeder roots compared to those at the heavy-textured site. Partially owning to the deeper and larger root system, H. ammodendron growing at coarse-textured site was in better water conditions than those at heavy-textured site under the same climatic conditions.     

Effects of simulated grazing on different genotypes of Bouteloua gracilis: how important is morphology?

Populations of grasses exposed to grazing by vertebrates often exhibit reduced stature, increased tillering, reduced flowering, and other morphological differences which distinguish them from ungrazed populations. These differences frequently are interpreted as an adaptive response that reduces grazing damage; however, there are few experimental tests of this hypothesis. This paper describes a field experiment designed to determine whether morphological variation among genotypes of the grass Bouteloua gracilis is related to variation in their responses to grazing. Eleven genotypes differing in morphological and reproductive characters were transplanted into a shortgrass steppe community near Fort Collins, Colorado. Replicates of each genotype were subjected to clipping treatments intended to realistically simulate three grazing intensities. After two growing seasons, different genotypes still maintained significant differences in a wide range of morphological and demographic characters. However, there were few significant effects of grazing treatment, and no significant genotype×treatment interactions. These results suggest that for B. gracilis clipped in simulation of natural grazing, defoliation has few short-term effects on fitness components, and intrapopulation morphological variation has few consequences for defoliation resistance. Received: 22 March 1999 / Accepted: 26 October 1999     

Response of grassland biomass production to simulated climate change and clipping along an elevation gradient

Changes in rainfall and temperature regimes are altering plant productivity in grasslands worldwide, and these climate change factors are likely to interact with grassland disturbances, particularly grazing. Understanding how plant production responds to both climate change and defoliation, and how this response varies among grassland types, is important for the long-term sustainability of grasslands. For 4 years, we manipulated temperature [ambient and increased using open-top chambers (OTC)], water (ambient, reduced using rainout shelters and increased using hand watering) and defoliation (clipped, and unclipped) in three grassland types along an elevation gradient. We monitored plant cover and biomass and found that OTC reduced biomass by 15 %, but clipping and water treatments interacted with each other and their effects varied in different grassland types. For example, total biomass did not decline in the higher elevation grasslands due to clipping, and water addition mitigated the effects of clipping on subordinate grasses in the lower grasslands. The response of total biomass was driven by dominant plant species while subordinate grasses and forbs showed more variable responses. Overall, our results demonstrate that biomass in the highest elevation grassland was least effected by the treatments and the response of biomass tended to be dependent on interactions between climate change treatments and defoliation. Together, the results suggest that ecosystem function of these grasslands under altered climate patterns will be dependent on site-specific management.     

Plant competition,abiotic, and long- and short-term effects of large herbivores on demography of opportunistic species in a semiarid grassland

Summary The emergence and subsequent survival and growth of five opportunistic weeds were monitored after seed additions to long-term grazing treatments with or without current-year grazing, long-term ungrazed treatments, and removal treatments designed to eliminate plant competition from existing perennials while either leaving vegetation and soil structure unaltered or disturbed. The treatments were applied on both uplands and lowlands to assess the relative influence of macroabiotic environment versus plant competition. The long-term effects of large herbivores on the initial emergence of seedlings were greater than the effects of removing competition. Very few individuals emerged on the long-term grazed treatments that were either grazed or ungrazed during the experiment. Numbers of individuals emerging on the long-term ungrazed treatments were greater or equal to those emerging on the no-competition-undisturbed treatments, but numbers were greatest on no-competition-disturbed treatments. None of the seeded individuals on the long-term grazed, currently grazed treatments survived to the end of the growing season. There was a slightly greater end-of-season biomass of seeded species and percentage of the total population reaching reproductive status on the long-term ungrazed compared with grazed-nondefoliated treatments, and very high survival, biomass, and proportions of reproductives on both no-competition treatments. Cover types in the immediate vicinity of seedlings influenced both germination and survival, but the effects differed between species and treatments. Equal compensation to current-year herbivory occurred on long-term heavily grazed treatments even though above-ground production was much greater on long-term protected sites. Productivity varied with topography, but very few topographic main effects or interactions occurred with demographic variables of seeded species, suggesting that macroabiotic effects were of minor importance compared with grazing and plant competition.     

Seed bank composition and above-ground vegetation in response to grazing in sub-Mediterranean oak forests (NW Greece)

We investigate the persistent soil seed bank composition and its relation to the above-ground flora of grazed and non-grazed sub-Mediterranean deciduous oak forests of NW Greece. Twenty-eight taxa were recorded in the soil seed bank and 83 taxa (70 taxa in plots of seed bank sampling) in the above-ground vegetation. The dominant tree species and many woodland species found in the above-ground vegetation were absent from the soil seed bank. Similarity between the soil seed bank and the above-ground vegetation decreased with grazing, and grazing led to a decrease of species richness in above-ground vegetation and soil seed bank. Beta diversity of vegetation among grazed and among non-grazed plots did not differ, but was significantly higher between grazed and non-grazed areas. Beta diversity of the soil seed bank declined with grazing. When applying classification tree and logistic regression analyses, non-grazed forest sites are clearly differentiated by the presence of Phillyrea latifolia, Euphorbia amygdaloides and Brachypodium sylvaticum. PCA ordination of above-ground species composition reflected a gradient from sites grazed by ruminants to non-grazed sites, but no clear structure was detected in the seed bank.     

How season of grazing and herbivore selectivity influence monsoon tall-grass communities of northern Australia

Abstract. The hypothesis that season of defoliation and herbivore selectivity may be as important as level of use in determining plant community response to grazing was tested in a monsoon grassland in northern Australia. Plots, dominated by the tussock grasses Themeda triandra and Chrysopogon fallax, were grazed by cattle at low, medium and high rates of utilization in either the early wet, late wet or dry seasons. Effects of grazing on species composition were greatest in the early wet season when high rates of utilization significantly reduced the proportion and occurrence of Themeda and increased the proportion of forbs. Grazing in the dry season had no significant effect on composition. At medium and high levels of utilization in the early wet season, the pasture responded negatively to defoliation, only partially compensating for plant tissue lost to herbivory. The negative response to defoliation carried over to the next wet season when these same medium and high-grazing treatments produced only 80 % and 60 % growth, respectively, of that in treatments grazed at low levels of utilization or those grazed during the dry season. The frequency of Themeda was still lower, and that of annual grasses and non-leguminous forbs higher, in plots that had been grazed at a high rate of utilization for just eight weeks in the early wet season two years previously. Species richness and diversity were also significantly affected by this grazing disturbance. If species composition is to be maintained in these grasslands then stocking rates must be set at low levels to cope with the combined effect of undercompensation in response to defoliation in the wet season and strong dietary preferences for grazing sensitive species.     

Impact of selective grazing on plant production and quality through floristic contrasts and current-year defoliation in a wet grassland

Grazing impacts the structure and functional properties of vegetation through floristic changes (i.e., long-term effect) and current defoliation (i.e., short-term effect). The aim of this study was to assess the relative importance of these two grazing effects on productivity (ANPP) and plant quality (C/N ratio) among plant patches submitted to a variety of grazing intensity for several years. Long-term grazing effect was measured by comparing ANPP and C/N ratio among plant patches with contrasting floristic composition. Short-term impact of grazing was measured by comparing ANPP and C/N in plant patches, with and without defoliation. Floristic contrasts led to a lower ANPP in highly grazed patches than in lightly grazed ones. This result may be related to the increasing proportion of grazing-tolerant and grazing-avoiding species with increasing grazing intensity. Vegetation C/N contrasts were recorded among grazed patches but did not linearly relate to grazing intensity. Short-term effect of current-year defoliation on ANPP was limited as vegetation compensated for biomass removal. No evidence for grazing-enhancement of ANPP was found even at moderate grazing intensity. Long-term floristic changes with grazing thus appeared to be the main driving factor of variations in ANPP. In contrast, C/N ratio showed no general and consistent variation along the grazing gradient but varied consistently depending on the community investigated, thus suggesting an effect of the species pool available.     

Morphologic and allozymic variation between long-term grazed and non-grazed populations of the bunchgrass Schizachyrium scoparium var. frequens

Summary Plant populations of Schizachyrium scoparium var. frequens with a history of long-term grazing by domestic herbivores were characterized by shorter and narrower leaf blades and tillers of lower weight than plants from populations with a history of no grazing. Following four biweekly defoliation events plants from the grazed populations additionally displayed lower specific leaf weights, lower amounts of biomass removed per tiller upon defoliation and a greater number of new and regrowing tillers. Production values per plant remained similar between the two populations because of a greater number of tillers per plant in the grazed population.A large amount of allozymic polymorphism was observed in both the grazed and non-grazed populations. Results of the electrophoretic analyses indicated that a minimum of 66% of the plants sampled represented separate genotypes. No distinction could be made between grazed and non-grazed populations in terms of allozymic partitioning. The morphological variation observed between these two populations may represent a shift in the relative dominance of a series of genotypes variously adapted to herbivory.     

Herbivory and ramet performance in the clonal herb Trientalis europaea L.

1 The intensity of leaf damage caused by invertebrate herbivores and grazing by vertebrates and their effects on clonal growth, survival and reproduction were examined in a population of the forest herb Trientalis europaea during 4 years.
2 Levels of herbivory were low and varied between years in the studied population. Larger ramets were exposed to a greater risk of both grazing and leaf defoliation. Ramets that suffered leaf damage in 1 year experienced an increased probability of leaf damage in succeeding years. The probability of suffering herbivore attack was independent among the ramets belonging to the same clonal fragment, suggesting that clonal propagation might operate as a risk-spreading strategy in this species.
3 Leaf damage did not affect any measure of plant performance, probably due to the low amounts of leaf area removed by invertebrate herbivores. In contrast, vertebrate grazing affected all phases of the pseudo-annual life cycle of T. europaea . Grazing prevented flowering and fruiting, increased ramet mortality during summer and decreased tuber production. Furthermore, grazed ramets produced shorter stolons and smaller tubers, which in turn had a lower winter survival and produced smaller ramets in the following growing season. The large impact of grazing was due to the consumption of the whole of the single shoot of ramets of T. europaea . Although regrowth was possible, secondary shoots were significantly smaller and assimilation was delayed.
4 Tubers originating from grazed ramets were placed shallower than tubers from ungrazed ramets. This could be due to an alteration in the growth pattern of stolons of the grazed ramets.     

Production and turnover rates of shallow fine roots in rangelands of the Patagonian Monte, Argentina

Selective sheep grazing in arid rangelands induces a decrease in total cover and grass cover and an increase in the dominance of shrubs. Both life forms differ in aboveground and belowground traits. We hypothesized that grazing disturbance leads to the replacement of grass by shrub fine roots in the upper soil, and this is reflected in changes in the seasonal dynamics of shallow fine roots at the community level. In two sites representative of non-grazed and grazed vegetation states in the Patagonian Monte, we assessed the canopy structure, and the fine root biomass, N concentration, production, and turnover during two consecutive years. The non-grazed site exhibited higher total, grass, and shrub cover than the grazed site. The grazed site had larger or equal fine root biomass than the non-grazed site except for late spring of the second year. This could be associated with the ability of shrubs to develop dimorphic-root systems occupying the soil freed by grasses at the grazed site, and with the larger contribution of grass than shrub fine roots in relation to an extraordinary precipitation event at the non-grazed site. This was consistent with the N concentration in fine roots. Fine root production was positively correlated to temperature at the grazed site and with precipitation at the non-grazed site. Fine root turnover did not differ between sites. Our results indicate that grazing leads to a shifting in the seasonality and main climatic controls of fine root production, while fine root turnover is mostly affected by changes in soil water conditions.