Competitive ability of the bunchgrass Schizachyrium scoparium as affected by grazing history and defoliation

Herbivory by large animals is known to function as a selection pressure to increase herbivory resistance within plant populations by decreasing the frequency of genotypes possessing large, erect canopies. However, the increase in herbivory resistance of the remaining genotypes in the population may potentially involve a tradeoff with competitive ability. The perennial bunchgrass Schizachyrium scoparium was grown in a transplant garden to test the hypothesis that late successional plant populations with a history of grazing are at a competitive disadvantage relative to conspecific populations with no history of grazing were found to possess a greater competitive ability than plants with no grazing history in the absence of herbivory. This unexpected response resulted from the capacity of plants with a history of grazing to recruit a greater number of smaller tillers than did plants with no grazing history. This response was only significant when plants with a history of grazing were nondefoliated and grown with the weakest of the mid-successional competitors, indicating that both defoliation and intense interspecific competition can mask the architectural expression of herbivore-induced selection. Individual tillers did not display any architectural differences between plants with contrasting grazing histories other than mean tiller weight. These data confirm that herbivory by domestic cattle may function as a selection pressure to induce architectural variation in grass populations within an ecological time frame (ca <-25 yrs).     

Does selective defoliation mediate competitive interactions in a semiarid savanna? A demographic evaluation

Abstract. Three patterns of target-neighbor plant defoliation were imposed on a late-seral, perennial, C₄-grass, Bouteloua curtipendula, in three long-term grazing regimes to determine the influence of selective defoliation on competitive interactions and species replacement in a semiarid savanna on the Edwards Plateau, Texas, USA. Short-term (3-yr) target plant defoliation did not significantly affect either tiller or plant responses in any of the three grazing regimes. Neighbor plant defoliation, either alone or in combination with target plants, produced a significant defoliation interaction with time for tiller number and basal area per plant, but not for tiller recruitment or mortality. The minimal effect of selective defoliation on the intensity of competitive interactions in this semiarid community indicates that selective grazing has a less definitive role in mediating herbivore-induced species replacement than it does in mesic grasslands and savannas. This interpretation is discussed within the context of long-term (45-yr) change in herbaceous vegetation associated with grazing in this community. Cumulative tiller recruitment in the intensively grazed regime was only 44% of that in the ungrazed regime because of greater plant mortality and fewer surviving plants that recruited tillers. Target plant mortality (50%) only occurred in the intensively grazed regime and the proportion of target plants that initiated tillers decreased by 70, 48 and 32% in the ungrazed, moderately and intensively grazed regimes, respectively, during the final two years of the investigation. The decrease in cumulative tiller recruitment in all grazing regimes was probably mediated by a drought-induced increase in median tiller age the second year of the study. However, tiller per tiller recruitment rate among plants that recruited at least one tiller remained relatively constant among grazing regimes and years. Intensive, long-term grazing has modified the population structure of this late-seral perennial grass to the extent that population responses to both herbivory and periodic drought have been altered in comparison with those of ungrazed and moderately grazed populations. Ecological consequences of a herbivore-induced transition in population structure may be to minimize the effect of selective herbivory on competitive interactions and to function as an avoidance mechanism to reduce the probability of localized population extinction in response to intensive long-term herbivory.     

Patterns in grass silicification: response to grazing history and defoliation

Summary Morphologically distinct populations of a North American perennial grass, Agropyron smithii, collected from a heavily grazed prairie dog (Cynomys ludovicianus) colony (PDC) and a grazing exclosure (EX), were grown in an environmental chamber to determine whether: (1) leaf silicon (Si) concentrations are greater in plant populations which differentiated under heavy grazing pressure, and (2) leaf silicification is inducible by defoliation. Mean shoot Si concentration of nondefoliated plants was greater in the PDC population (2.2%) than the EX population (1.9%) over the 18 wk experiment, largely as a result of differences in Si concentrations in leaf blades. However, leaf Si concentration was lower in defoliated plants of each population than in nondefoliated plants, indicating that leaf silicification was not an inducible herbivore defense mechanism in A. smithii. The higher leaf Si concentrations from the heavily grazed population may be associated with grazingrelated environmental stresses such as a warmer, drier microclimate or with morphological characteristics related to grazing tolerance or avoidance.     

Genetic adaptations to grazing and mowing in the unpalatable grass Cenchrus incertus

Summary To identify morphological and life history adaptations to grazing, mowing, and cultivation, seeds of the grass Cenchrus incertus were collected from two populations in each of three types of sites: cemeteries (mown occasionally), pastures (grazed continuously), and orchards (plowed twice a year). Seeds from each population were germinated and grown in a common greenhouse.Plants originating from the two cemetery populations had, on average, the most leaves and the most tillers per plant at each census, and they were on average the shortest in stature. Cemetery plants had on average the greatest number of panicles and of burs per plant, but the fewest burs per panicle. The occasionally-mown but ungrazed cemetery populations in this study were therefore more similar to grazed populations described in other studies; the pasture and orchard populations in this study were more similar to ungrazed populations described in other studies. We suggest that this may be due to the low acceptability of Cenchrus incertus, which makes its defoliation relatively infrequent in unmown sites.Some of the traits that distinguished the cemetery populations from the orchard and pasture populations, such as shorter stature, are probably direct adaptations to defoliation. Others may be secondary effects of these, or the result of allocation trade-offs.     

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.     

Soil and plant water relations in a crested wheatgrass pasture: response to spring grazing by cattle

Summary Few field studies have attempted to relate effects of actual livestock grazing on soil and plant water status. The present study was initiated to determine the effects of periodic defoliations by cattle during spring on soil moisture and plant water status in a crested wheatgrass (Agropyron cristatum (L.) Gaertn. and A. desertorum (Fisch. ex Link) Schult.) pasture in central Utah. Soil moisture in the top 130 cm of the soil profile was depleted more rapidly in ungrazed plots than in grazed plots during spring and early summer. Soil moisture depletion was more rapid in grazed plots in one paddock after 1 July due to differential regrowth, but there was no difference in soil water depletion between plots in another paddock during the same period. This difference in soil water depletion between paddocks was related to a difference in date of grazing. Although more water had been extracted from the 60 cm to 130 cm depths in ungrazed plots by late September, cumulative soil moisture depletion over the entire 193 cm profile was similar in grazed and ungrazed plots. Prior to 1 July, grazing had no effect on predawn leaf water potentials as estimated by a pressure chamber technique; however, after 1 July, predawn leaf water potentials were lower for ungrazed plants. Midday leaf water potentials were lower for grazed plants before 1 July, but did not differ between grazed and ungrazed plants after 1 July. A 4- to 8-day difference in date of defoliation did not affect either predawn or midday leaf water potentials. The observed differences in water use patterns during spring and early-summer may be important in influencing growth and competitive interactions in crested wheatgrass communities that are subject to grazing by domestic livestock.     

Variation in grazing tolerance among three tallgrass prairie plant species

Three tallgrass prairie plant species, two common perennial forbs (Artemisia ludoviciana and Aster ericoides [Asteraceae]) and a dominant C(4) perennial grass (Sorghastrum nutans) were studied under field and greenhouse conditions to evaluate interspecific variation in grazing tolerance (compensatory growth capacity). Adaptation to ungulate grazing was also assessed by comparing defoliation responses of plants from populations with a 25-yr history of no grazing or moderate ungulate grazing. Under field conditions, all three species showed significant reductions in shoot relative growth rates (RGR), biomass, and reproduction with defoliation. In the two forbs, clipping resulted in negative shoot RGR and reductions in both number and length of shoot branches per ramet. Sorghastrum nutans maintained positive RGR under defoliation due to a compensatory increase in leaf production. Defoliation reduced rhizome production in A. ericoides and S. nutans, but not in A. ludoviciana. Clipping significantly reduced sexual reproductive allocation in all three species, although S. nutans showed a smaller reduction than the forbs. All three species showed similar responses to defoliation in burned and unburned sites. Under greenhouse conditions, a similar clipping regimen resulted in smaller reductions in growth and reproduction than those observed in the field. For all three species, the grazing tolerance indices calculated under natural field conditions were significantly lower than those estimated from greenhouse-grown plants, and the interspecific patterns of grazing tolerance were different. Aster ericoides exhibited the highest overall defoliation tolerance under greenhouse conditions, followed by S. nutans. Artemisia ludoviciana, the only study species that is typically not grazed by ungulates in the field, showed the lowest grazing tolerance. In the field experiment S. nutans showed the highest grazing tolerance and the two forbs had similar low tolerance indices. These patterns indicate that, despite high compensatory growth potential, limited resource availability and competition in the field significantly reduce the degree of compensation and alter interspecific differences in grazing tolerance among prairie plants. In all three species, defoliation suppressed sexual reproduction more than growth or vegetative reproduction. Significant interactions between plant responses to defoliation and site of origin (historically grazed or ungrazed sites) for some response variables (root/shoot ratios, rhizome bud initiation, and reproductive allocation) indicated some degree of population differentiation and genetic adaptation in response to a relatively short history of ungulate grazing pressure. The results of this study indicate that patterns of grazing tolerance in tallgrass prairie are both genetically based and also environmentally dependent.     

Coping with herbivory: Photosynthetic capacity and resource allocation in two semiarid Agropyron bunchgrasses

Summary Agropyron desertorum, a grazing-tolerant bunchgrass introduced to the western U.S. from Eurasia, and Agropyron spicatum, a grazing-sensitive bunchgrass native to North America, were examined in the field for photosynthetic capacity, growth, resource allocation, and tiller dynamics. These observations allowed identification of physiological characteristics that may contribute to grazing tolerance in semiarid environments. A uniform matrix of sagebrush, Artemisia tridentata, provided an ecologically relevant competitive environment for both bunch-grass species. Physiological activity, growth, and allocation were also followed during recovery from a severe defoliation treatment and were correlated with tiller dynamics.Potential photosynthetic carbon uptake of both species was dominated by stems and leaf sheaths during June, when maximum uptake rates occurred. For both species, water use efficiency of stems and sheaths was similar to that of leaf blades, but nitrogen investment per photosynthetic surface area was less than in blades. In addition, soluble carbohydrates in stems and sheaths of both species constituted the major labile carbon pools in control plants. Contrary to current theory, these findings suggest that culms from which leaf blades have been removed should be of considerable value to defoliated bunchgrasses, and in the case of partial defoliation could provide important supplies of organic nutrients for regrowth. These interpretations, based on total pool sizes, differ markedly from previous interpretations based on carbohydrate concentrations alone, which suggested that crowns contain large carbohydrate reserves. In this study, crowns of both species contained a minor component of the total plant carbohydrate pool.Following defoliation, A. desertorum plants rapidly reestablished a canopy with 3 to 5 times the photosynthetic surface of A. spicatum plants. This difference was primarily due to the greater number of quickly growing new tillers produced following defoliation. Agropyron spicatum produced few new tillers following defoliation despite adequate moisture, and carbohydrate pools that were equivalent to those in A. desertorum.Leaf blades of regrowing tillers had higher photosynthetic capacity than blades on unclipped plants of both species, but the relative increase, considered on a unit mass, area, or nitrogen basis, was greater for A. desertorum than for A. spicatum. Agropyron desertorum also had lower investment of nitrogen and biomass per unit area of photosynthetic tissues, more tillers and leaves per bunch, and shorter lived stems, all of which can contribute to greater tolerance of partial defoliation.Greater flexibility of resource allocation following defoliation was demonstrated by A. desertorum for both nitrogen and carbohydrates. Relatively more allocation to the shoot system and curtailed root growth in A. desertorum resulted in more rapid approach to the preclipping balance between the root and shoot systems, whereas root growth in A. spicatum continued unabated following defoliation. Nitrogen required for regrowth in both species was apparently supplied by uptake rather than reserve depletion. Carbohydrate pools in the shoot system of both species remained very low following severe defoliation and were approximately equivalent to carbon fixed in one day by photosynthesis of the whole canopy.Dedicated to Drs. Michael Evenari and Konrad Springer     

GRAZING HISTORY,DEFOLIATION, AND FREQUENCY-DEPENDENT COMPETITION: EFFECTS ON TWO NORTH AMERICAN GRASSES

Agropyron smithii and Bouteloua gracilis plants from intensively grazed prairie dog colonies and from a grazing exclosure in Wind Cave National Park, South Dakota, were used to compare responses of conspecific populations with different histories of exposure to grazing and to competition for light. In separate experiments for each species, plants grown in monocultures and two-population replacement-series mixtures were used to examine effects of defoliation, frequency-dependent competition, and population on biomass and morphology. Colony and exclosure plants frequently responded differently. Defoliation more often adversely affected exclosure plants than colony plants, while interpopulation competition more often adversely affected colony plants. Defoliation frequently negated the competitive advantage of exclosure plants. Intrapopulation competition appeared to be greater among exclosure than colony plants. Our results indicate that conclusions based on studies of plants in long-term exclosures may not apply to plant populations having long histories of intensive grazing. While there were differences between species, in both, these experiments provide evidence of population differentiation, resulting in morphologically dissimilar populations which responded differently to defoliation and to inter- and intrapopulation competition.     

Response of galling invertebrates on Salix lanata to reindeer herbivory

Browsing and defoliation often increase the densities of insect herbivores on woody plants. Densities of herbivorous invertebrates were estimated in a long-term grazing manipulation experiment. More then 30-yr-old fences allow us to compare densities of invertebrate herbivores on Salix lanata in areas heavily grazed and areas lightly grazed by reindeer. The number of gall-forming insects ( Pontania glabrifons) and gall-forming mites were higher on the heavily grazed shrubs than on lightly grazed shrubs. In contrast to most short-term studies, the heavily grazed S. lanata had shorter current annual shoots. No difference in leaf size, leaf nitrogen content, or C:N ratio between grazing intensities were detected. However, the enhanced natural δ15N value indicates that heavily grazed shrubs get a higher proportion of their N directly from reindeer faeces. Leaf weight per unit area and relative fluctuating asymmetry of leaf shape increased in heavily grazed S. lanata . Enhanced relative fluctuating asymmetry might indicate higher susceptibility to herbivores. Long-term grazing seems to increase the density of invertebrate herbivory in the same way as short-term grazing, even if the plant responses differ substantially.     

The ten year cycle of the willow grouse of Lower Kolyma

Summary The effects of defoliation on growth and nitrogen (N) nutrition were examined in populations of Agropyron smithii (western wheatgrass) collected from a heavily grazed black-tailed prairie dog (Cynomys ludovicianus) colony (ON-colony) and a nearby lightly grazed, uncolonized area (OFF-colony). Defoliated and nondefoliated plants were grown at low soil N availability with similar sized defoliated individuals of A. smithii from a grazing-exclosure population as a common competitor. Sequential harvests were made over 24 days following defoliation. Growth analysis plus biomass and N yield and distribution data were used to identify features which may contribute to plant defoliation tolerance. Defoliation reduced total production 34% across populations. Defoliated plants produced as much new blade tissue, but only 67% as much new root biomass as did nondefoliated controls. Plants from prairie dog colonies accumulated biomass at a faster relative rate than did plants from uncolonized sites, in part, because of a 250% greater mean relative growth rate of blades and more than 200% greater rate of biomass production per unit blade biomass. Total N accumulation was significantly greater in defoliated ON- than OFF-colony individuals. The mean relative accumulation rate of N was increased by defoliation in ON-colony plants, but reduced by defoliation in OFF-colony plants. The mean rate of N accumulation per unit root biomass was more than 300% greater in the ON- than OFF-colony population. Colony plants initially had a greater proportion of biomass and N remaining after defoliation in roots. Initial differences between populations in the distribution of biomass and N were eliminated as colony plants concentrated 24-day accumulation of biomass and N in aboveground structures. The data suggest that the combination of growth, N nutrition, and biomass and N distribution characteristics of the colony population likely confer a high rate of resource capture on heavily grazed prairie dog colonies.     

Performance of two Picea abies (L.) Karst. stands at different stages of decline

Summary The annual replacement of tillers of Agropyron desertorum (Fisch. ex Link) Schult., a grazing-tolerant, Eurasian tussock grass, was examined in the field following cattle grazing. Heavy grazing before internode (culm) elongation seldom affected tiller replacement. Heavy grazing during or after internode elongation, which elevates apical meristems, increased overwinter mortality of fall-produced tillers and reduced the number and heights of these replacement tillers. Unexpectedly, tussocks grazed twice within the spring growing season tended to have lower overwinter tiller mortality, greater tiller replacement, and larger replacement tillers than tussocks grazed only once in late spring. These responses of twice-grazed tussocks, however, were still less than those of ungrazed tussocks or tussocks grazed moderately in early spring. The presence of ungrazed tillers on partially grazed tussoks did not increase the replacement of associated grazed tillers relative to tillers on uniformly grazed plants. This result indicates that resource sharing among tillers, if present, is short-lived or ecologically unimportant in this species. Although A. desertorum is considered grazing-tolerant, tiller replacement on heavily grazed tussocks, particularly those grazed during or after internode elongation when apical meristems were removed, was usually inadequate for tussock maintenance. These observations at the tiller (ramet) level of organization in individual tussocks (genet) may explain the often noted reduction in stand (population) longevity with consistent heavy grazing.     

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

Summary The 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.     

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.     

Axillary bud banks of two semiarid perennial grasses: occurrence, longevity, and contribution to population persistence

The occurrence, longevity, and contribution of axillary bud banks to population maintenance were investigated in a late-seral perennial grass, Bouteloua curtipendula, and a mid-seral perennial grass, Hilaria belangeri, in a semiarid oak-juniper savanna. Axillary buds of both species were evaluated over a 2-year period in communities with contrasting histories of grazing by domestic herbivores. A double staining procedure utilizing triphenyl tetrazolium chloride and Evan's blue indicated that both viable and dormant axillary buds remained attached to the base of reproductive parental tillers for 18–24 months which exceeded parental tiller longevity by approximately 12 months. Bud longevity of the late-seral species, B. curtipendula, exceeded bud longevity of the mid-seral species, H. belangeri, by approximately 6 months. Younger buds located on the distal portion of the tiller base were 3.2 and 1.4 times more likely to grow out than older proximal buds of B. curtipendula and H. belangeri, respectively. The percentage of older proximal buds, which included comparable portions of viable and dormant buds, that grew out to produce tillers following mortality of parental tillers was 6.0% for B. curtipendula and 8.4% for H. belangeri. In spite of the occurrence of relative large axillary bud banks for both species, the magnitude of proximal bud growth did not appear sufficient to maintain viable tiller populations. We found no evidence to support the hypothesis of compensatory bud growth on an individual tiller basis for either species. Grazing history of the communities from which the buds were collected did not substantially affect the number, status, longevity, or outgrowth of axillary buds on an individual tiller basis for either species. However, long-term grazing by domestic herbivores influenced axillary bud availability by modifying population structure of these two species. Bud number per square meter for B. curtipendula was 25% lower in the long-term grazed compared to the long-term ungrazed community based on a reduction in both tiller number per plant and plant number per square meter. In contrast, bud number per square meter for H. belangeri was 190% greater in the long-term grazed than in the long-term ungrazed community based on a large increase in plant density per square meter. Minimal contributions of axillary bud banks to annual maintenance of tiller populations in this mid- and late-seral species underscores the ecological importance of consistent tiller recruitment from recently developed axillary buds. Consistent tiller recruitment in grasslands and savannas characterized by intensive grazing and periodic drought implies that (1) bud differentiation and maturation must be remarkably tolerant of adverse environmental conditions and/or (2) tiller recruitment may resume from buds that mature following the cessation of severe drought and/or grazing, rather than from mature buds that survive these disturbances. These scenarios warrant additional research emphasis given the critical importance of this demographic process to tiller replacement in species populations and the maintenance of relative species abundance in grasslands and savannas. Received: 12 August 1996 / Accepted: 30 December 1996     

The impact of grazing intensity on soil characteristics of Stipa grandis and Stipa bungeana steppe in northern China (autonomous region of Ningxia)

The effects of grazing intensity on selected soil characteristics in the feather-grass steppes of the autonomous region of Ningxia (northern China) were investigated by a comparison of non-grazed areas (grazing intensity 0), slightly grazed areas (grazing intensity I), moderately grazed areas (II), intensively grazed areas (III) and over-grazed areas (IV). Even in areas used only minimally for grazing activities (I), a serious increase (doubling) in soil hardness was apparent in the upper soil layer. A continual decrease in organic matter in the surface soil can be correlated directly to soil compaction. The content of organic matter in soil of degree IV amounts to only a third of the organic matter found in non-grazed areas. This decrease can be attributed partly to the poor living conditions for soil organisms in compacted soils, but also to a significant reduction in litter. This is because intensive grazing causes reduced vegetation cover leading to litter being blown away by wind or washed away by heavy rainfall. Thus in level III hardly any plant litter remained to be incorporated into the soil as humus. Likewise root density also suffered its largest decrease in areas with a grazing intensity level III. With regard to the content of nitrogen and phosphorous (total and available) hardly any difference between soils of grazing intensity 0 and I was observed, whereas a noticeable decrease was apparent between levels I and II. Available Potassium was similar for all grazing levels. The pH-value of the soil solution is not significantly affected by grazing. We did not observe differences in the soils of the two main types of steppe vegetation (Stipa grandis and Stipa bungeana steppe) in response to grazing. Only the amount of litter in the S. grandis-steppe in non-grazed or slightly grazed areas is noticeably higher than in the S. bungeana steppe.     

Demography of three dominant sedges under contrasting grazing regimes in the High Arctic

Abstract. Tiller demography of Carex aquatilis ssp. stans, Carex membranacea, and Eriophorum angustifolium ssp. triste was investigated in ungrazed and grazed high arctic vegetation on central Ellesmere Island, Canada. Tiller birth, growth, flowering and death were studied from excavated clonal fragments, and tiller density and biomass were studied from excavated turfs. Five life‐cycle stages were determined: dormant buds, juvenile, mature, flowering and dead tillers. A stage‐based transition matrix model was developed to estimate the long‐term dynamics of the sedge populations and to compare life‐history strategies between ungrazed and grazed populations. Short‐term and retrospective models, based on the growth during the sampling year and during the lifetime of the clonal fragments, respectively, were compared to see how well the short‐term model can describe demography of long‐lived plants. According to the short‐term model, tiller populations were decreasing (λ < 1 except for C. membranacea), whereas the retrospective model indicated that the tiller populations were increasing. Tiller population growth rates did not differ between ungrazed and grazed habitats. Nevertheless, the similar growth rates may be obtained by balanced differences in the vital rates between plants of the two habitats. The plants in the ungrazed habitat tended to remain in their current life‐cycle stage, whereas plants in the grazed habitat moved quickly to the next stage and died earlier. C. aquatilis ssp. stans appears to gain a competitive advantage over the other species under intensive grazing, as indicated by the higher tiller density and greater below‐ground biomass in grazed vegetation. The greater amount of below‐ground biomass apparently buffers C. aquatilis ssp. stans against grazing better than the other species.     

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.     

Defoliation of a grass is mediated by the positive effect of dung deposition,moss removal and enhanced soil nutrient contents: results from a reindeer grazing simulation experiment

Herbivory is one of the key drivers shaping plant community dynamics. Herbivores can strongly influence plant productivity directly through defoliation and the return of nutrients in the form of dung and urine, but also indirectly by reducing the abundance of neighbouring plants and inducing changes in soil processes. However, the relative importance of these processes is poorly understood. We, therefore, established a common garden experiment to study plant responses to defoliation, dung addition, moss cover, and the soil legacy of reindeer grazing. We used an arctic tundra grazed by reindeer as our study system, and Festuca ovina, a common grazing‐tolerant grass species as the model species. The soil legacy of reindeer grazing had the strongest effect on plants, and resulted in higher growth in soils originating from previously heavily‐grazed sites. Defoliation also had a strong effect and reduced shoot and root growth and nutrient uptake. Plants did not fully compensate for the tissue lost due to defoliation, even when nutrient availability was high. In contrast, defoliation enhanced plant nitrogen concentrations. Dung addition increased plant production, nitrogen concentrations and nutrient uptake, although the effect was fairly small. Mosses also had a positive effect on aboveground plant production as long as the plants were not defoliated. The presence of a thick moss layer reduced plant growth following defoliation. This study demonstrates that grasses, even though they suffer from defoliation, can tolerate high densities of herbivores when all aspects of herbivores on ecosystems are taken into account. Our results further show that the positive effect of herbivores on plant growth via changes in soil properties is essential for plants to cope with a high grazing pressure. The strong effect of the soil legacy of reindeer grazing reveals that herbivores can have long‐lasting effects on plant productivity and ecosystem functioning after grazing has ceased.     

Comparative responses of the Savanna grasses Cenchrus ciliaris and Themeda triandra to defoliation

Summary Two perennial tussock grasses of savannas were compared in a glasshouse study to determine why they differed in their ability to withstand frequent, heavy grazing; Cenchrus ciliaris is tolerant and Themeda triandra is intolerant of heavy grazing. Frequent defoliation at weekly intervals for six weeks reduced shoot biomass production over a subsequent 42 day regrowth period compared with previously undefoliated plants (infrequent) in T. triandra, but not in C. ciliaris. Leaf area of T. triandra expanded rapidly following defoliation but high initial relative growth rates of shoots were not sustained after 14 days of regrowth because of reducing light utilising efficiency of leaves. Frequently defoliated plants were slower in rate of leaf area expansion and this was associated with reduced photosynthetic capacity of newly formed leaves, lower allocation of photosynthate to leaves but not lower tiller numbers. T. triandra appears well adapted to a regime where defoliation is sufficiently infrequent to allow carbon to be fixed to replace that used in initial leaf area expansion. In contrast, C. ciliaris is better adapted to frequent defoliation than is T. triandra, because horizontally orientated nodal tillers are produced below the defoliation level. This morphological adaptation resulted in a 10-fold higher leaf area remaining after defoliation compared with similarly defoliated T. triandra, which together with the maintenance of moderate levels of light utilising efficiency, contributed to the higher leaf area and shoot weight throughout the regrowth period.