Effects of sheep grazing and temporal variability on population dynamics of the clonal herb <Emphasis Type="Italic">Geranium sylvaticum</Emphasis> in an alpine habitat

An improved understanding of population-level consequences of grazing on plants can be facilitated by an assessment of grazing effects on all stages in the life-cycle. In this study, 6 years of demographic data for three populations of the perennial herb Geranium sylvaticum were analysed. We examined the effects of sheep grazing (high sheep density, low sheep density and no sheep) and interannual climatic variability on vital rates and population growth rates (λ). Grazing did not affect survival or flowering rates, but reduced rates of growth and increased rates of clonal reproduction. At the population level, high contributions from retrogression and clonal reproduction buffered reduced rates of growth and stasis, and no consistent differences in λ between populations exposed to different sheep densities were found. Instead, large between-year variability in λ, independent of sheep density, was detected, related to variation in the local summer climate. The results indicated, however, that grazing effects on λ were more severe in unfavourable than in normal years. Our study highlights that increased clonal reproduction rates functioned as a tolerance mechanism towards grazing in this herb, which forms a mechanism to explain how moderate population responses to grazing in some herbs can arise.     

Linking sheep density and grazing frequency to persistence of herb species in an alpine environment

Large herbivores are important drivers in ecosystems worldwide. Changes in herbivore densities are predicted to especially affect herbs that are strongly preferred by herbivores. The persistence of herbs could be challenged by enhanced grazing, but also grazing cessation may affect persistence, especially for prostrate herbs, which might be out-competed. To test how different herbivore densities (high, low, and no sheep) affect grazing frequency and plant responses (plant height, flowering frequency, and plant density) at the herb community and species level, we conducted a fully replicated, landscape-scale experiment in an alpine environment. We found that none of the herb species changed their densities after 5 years with experimental changes in grazing pressure. Sheep density affected grazing and flowering frequency at the herb community level. Eight herb species were more grazed at high sheep density as compared to enclosures with no sheep. Herb height decreased at high sheep density as compared to no sheep for the two species with the highest grazing frequency. Increased height and flowering frequency were found for small herbs at high sheep density. Our experiment clearly shows that herbs do not constitute a homogeneous functional group and that, in particular, tall and small herbs are affected in contrasting ways but all species (n = 15) tolerated changes in grazing regimes as densities were maintained at both enhanced grazing and grazing cessation.     

Insect herbivory and vertebrate grazing impact food limitation and grasshopper populations during a severe outbreak

1. Competition between herbivores often plays an important role in population ecology and appears strongest when densities are high or plant production is low. Phytophagous insects are often highly abundant, but relatively few experiments have examined competition between vertebrates and phytophagous insects. 2. In grassland systems worldwide, grasshoppers are often the dominant phytophagous insect, and livestock grazing is a dominant land use. For this study, a novel experiment was conducted examining competition between vertebrates and invertebrates, where both grasshopper densities and sheep grazing were manipulated inside 10‐m² caged mesocosms during a grasshopper outbreak. We examined how grasshopper densities and the timing of vertebrate herbivory affected grasshopper densities, if the effects of vertebrates on survival and reproduction changed with grasshopper density, and how a naturally occurring grasshopper outbreak affected grasshopper populations in the following year. 3. Densities of grasshoppers at the site peaked at 130 m^–2. Food‐limited competition was stronger in treatments with higher grasshopper densities and repeated or late livestock herbivory, leading to reduced survival, femur length, and functional ovarioles, a measure of future reproduction. Strong food‐limited density‐dependent reproduction and survival led to reduced hatching densities in 2001. 4. As competition was typically stronger with high grasshopper densities than with livestock grazing, competition from vertebrates could be relatively less important for phytophagous insect population dynamics during outbreaks. The experiment provides insights into how competition between insect and vertebrate herbivores influences insect population dynamics, and indicates that severe outbreaks can rapidly subside with strong competition from vertebrate and insect herbivores.     

Differential effects of herbivory and pathogen infestation on plant population dynamics

Plants often interact with antagonists such as herbivores or pathogens. Negative effects on individual plant performance are widely documented, but less is known about whether such effects translate into effects on population viability. In temperate forests, important herbivores include deer. During 2006–2009, I compared vital rates and population growth rates (calculated using integral projection models) between fenced exclosures and grazed control areas, using the perennial herb Phyteuma spicatum as a model species. Deer caused the largest damage to flowering individuals, removing about 24% of all inflorescences and 13% of the above-ground biomass. Only few vital rates seemed to be negatively affected by deer (mainly seed production) and this did not translate into effects on population growth rate. Contrary to expectations, population growth rates tended to be lower in the fenced exclosures in 1 year. This was likely caused by high-pathogen infestation rates, which negatively affected the probability of adult survival and growth. Population growth rate was more sensitive to changes in these vital rates than to changes in seed production. In summary, the results of this demographic study show that grazing effects may be small for long-lived herbs, and that negative effects on vital rates such as seed production may not always translate into effects on population growth rate. The findings also illustrate that other antagonists such as pathogens may be of greater relative importance for differences in population performance than herbivores.     

Demography of a dormancy-prone geophyte: influence of spatial scale on interpretation of dynamics

Both temporal and spatial scales are important in the evaluation of population dynamics, but the latter often receives less attention in demographic analyses. We used a 5-year demographic data set for a long-lived geophyte, Calochortus lyallii, to investigate the pattern and components of spatial variation at two scales (population and microsite). We found that neither the projected population structure nor asymptotic population growth rate (λ) varied greatly across either scale, although the underlying contributors to the variation in λ, V(λ), did differ between scales. Life table response experiment analyses showed that V(λ) among populations came primarily from variation in seedling survival and progression of non-reproductive plants, whereas V(λ) among microsites was primarily due to the variable fertility of large adults. Prolonged dormancy was important in reducing V(λ) among quadrats at both the scales, partly countering fluctuations in other transitions such as recruitment. This result represents some of the first evidence that underground “bulb banks” could function to offset the effects of a spatially heterogeneous environment in a manner analogous to seed banks. Future work is needed to isolate the specific, sometimes idiosyncratic, life history phenomena acting to modulate plant population dynamics in a spatial context.     

Remnant population dynamics in the facultative biennial Carum carvi in fragmented semi-natural grasslands

Transition matrix models were used to examine the population dynamics in the facultative biennial Carum carvi L. in semi-natural grasslands, specifically to assess what life cycle stages are important for population development and to evaluate the effects of environmental stochasticity on population persistence and, hence, the ability to develop remnant populations. The demographic studies were conducted over a 4-year period in three moderately grazed grasslands that differed in onset and duration of grazing. Experimental seed-sowing was also conducted in disturbed and undisturbed plots in the populations. Deterministic and stochastic models yielded overall negative population growth (λ < 1) for the populations. λ was sensitive to transitions in the most frequent vegetative stage classes. Elasticity analysis indicated that a large proportion of population growth could be ascribed to the stasis of individuals in the largest vegetative stage class. Life-table response experiment (LTRE) analyses showed also that progression to larger stage classes was important in explaining the between-population variation in λ. The expected time to extinction was on the order of several decades for the study populations. Seed-sowing indicated that seedling establishment was limited by both seed and micro-site availability. The populations of C. carvi seem to be able to persist for a rather long time in moderately grazed semi-natural grasslands, even in cases where populations are destined to become extinct. The results, thus, indicate that “biennials” are able to maintain remnant populations in managed semi-natural grasslands.     

Grazing responses in herbs in relation to herbivore selectivity and plant traits in an alpine ecosystem

Herbivores shape plant communities through selective foraging. However, both herbivore selectivity and the plant’s ability to tolerate or resist herbivory may depend on the density of herbivores. In an alpine ecosystem with a long history of grazing, plants are expected to respond to both enhanced and reduced grazing pressures, and the interaction between plant traits and changes in species abundance are expected to differ between the two types of alteration of grazing regime. To understand the mechanisms behind species response, we investigated the relationship between sheep selectivity (measured in situ), plant traits and experimentally derived measures of change in species abundance as a response to the enhancement (from low to high density) or cessation (from low to zero density) of sheep grazing pressure over a six-year time period for 22 abundant herb species in an alpine habitat in south Norway. Sheep selected large, late-flowering herbs with a low leaf C/N ratio. Species that increased in abundance in response to enhanced grazing pressure were generally small and had high root/shoot ratios, thus exhibiting traits that reflect both resistance (through avoidance) and tolerance (through regrowth capacity) strategies. The abundance of selected species remained stable during the study period, and also under the enhanced grazing pressure treatment. There was, however, a tendency for selected species to respond positively to cessation of grazing, although overall responses to cessation of grazing were much less pronounced than responses to enhanced grazing. Avoidance through short stature (probably associated with increased light availability through the removal of tall competitors) as well as a certain amount of regrowth capacity appear to be the main mechanisms behind a positive response to enhanced grazing pressure in this study. The plant trait perspective clearly improves our insight into the mechanisms behind observed changes in species abundance when the disturbance regime is altered. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

ENVIRONMENTAL AND GENETIC DETERMINANTS OF PLANT SIZE IN VIOLA SORORIA

Survivorship and fecundity in the forest herb, Viola sororia, are size-dependent. The basis of size variation among individuals of Viola sororia was investigated with a uniform environment experiment. Plants collected from natural populations were vegetatively reproduced and grown under two light regimes in a greenhouse. Analysis of quantitative variation showed: 1) significant differences between light treatments for characters related to plant shape and relative growth rate; 2) significant among-genet variation for plant size, plant shape and relative growth rate but none for physiological characters; and 3) a size threshold for cleistogamous seed production and rhizome production. Heritability estimates for the characters associated with plant size and shape ranged from 0.09 to 0.39, indicating significant genetic determination for these traits. In addition, among-genet differences in relative growth rate were substantial. The results of this study suggest that the size variation found in natural populations is not solely a function of environmental heterogeneity but is significantly influenced by the genotypes composing the population.     

Experimental evidence that livestock grazing intensity affects the activity of a generalist predator

Grazing by domestic ungulates has substantial impacts on ecosystem structure and composition. In grasslands of the northern hemisphere, livestock grazing limits populations of small mammals, which are a main food source for a variety of vertebrate predators. However, no experimental studies have described the impact of livestock grazing on vertebrate predators. We experimentally manipulated sheep and cattle grazing intensity in the Scottish uplands to test its impact on a relatively abundant small mammal, the field vole (Microtus agrestis), and its archetypal generalist predator, the red fox (Vulpes vulpes). We demonstrate that ungulate grazing had a strong consistent negative impact on both vole densities and indices of fox activity. Ungulate grazing did not substantially affect the relationship between fox activity and vole densities. However, the data suggested that, as grazing intensity increased i) fox activity indices tended to be higher when vole densities were low, and ii) the relationship between fox activity and vole density was weaker. All these patterns are surprising given the relative small scale of our experiment compared to large red fox territories in upland habitats of Britain, and suggest that domestic grazing intensity causes a strong response in the activity of generalist predators important for their conservation in grassland ecosystems.     

Experimental evidence that livestock grazing intensity affects cyclic vole population regulation processes

Grazing by domestic ungulates may limit the densities of small herbivorous mammals that act as key prey in ecosystems. Whether this also influences density dependence and the regulation of small herbivore populations, hence their propensity to exhibit multi-annual population cycles, is unknown. Here, we combine time series analysis with a large-scale grazing experiment on upland grasslands to examine the effects of livestock grazing intensity on the population dynamics of field voles (Microtus agrestis). Using log-linear modelling of replicated time series under different grazing treatments, we show that increased sheep densities weaken delayed density dependent regulation of vole population growth, hence reducing the cyclicity in vole population dynamics. While population regulation is commonly attributed to both top-down and bottom up processes, our results suggest that regulation of cyclic vole populations can be disrupted by the influence of another grazer in the same trophic level. These results support the view that ongoing changes in domestic grazing intensity, by affecting small mammal dynamics, can potentially have cascading impacts on higher trophic levels, and strongly influence the dynamics of upland grassland systems.     

Effects of small mammals and vertebrate predators on vegetation in the Chilean semiarid zone

 We monitored the cover and seed bank response of shrubs, perennial herbs, and ephemeral plants to experimental exclusion of both the principal rodent herbivore, Octodon degus, and its vertebrate predators from 1989 to 1994 in a semiarid Chilean mediterranean site. Although both richness and species composition of the plant community at the study site were largely determined by abiotic factors (mainly rainfall and soil nutrients), predator and herbivore exclusion had significant effects on the relative abundance of several plant species. Experimental exclusion of herbivores was associated with increased cover of some shrubs and a perennial grass, and decreased cover and seed densities of several ephemerals, especially those exotic or restricted to areas underneath shrubs. Herbivores apparently reduced shrubs through browsing and indirectly affected herb cover and seed densities by opening up areas under shrubs and/or modifying physical and chemical conditions of the soil. Plant responses to predator exclusion were less clear. Nevertheless, higher cover of some shrubs and ephemerals in the presence of predators suggests tritrophic effects through changes in small mammal densities and/or foraging behavior. Received: 22 April 1996 / Accepted: 14 August 1996     

Impacts of simulated livestock grazing on Utah prairie dogs (Cynomys parvidens) in a low productivity ecosystem

Allometric foraging theory suggests that herbivores of greatly differing size should co-exist through niche segregation, but a few studies of large–small herbivore foraging relationships have reported competitive interactions. This study addresses the potential roles of habitat productivity and large herbivore grazing intensities on large–small herbivore foraging interactions. We examined effects of different intensity simulated grazing treatments on forage abundance and quality for Utah prairie dogs (Cynomys parvidens) in a low productivity ecosystem, and consequent effects on prairie dog individual growth rates, foraging preferences, and activity budgets. We hypothesized that simulated grazing would have predominantly facilitative impacts on Utah prairie dogs, as was found for black-tailed prairie dogs in higher productivity ecosystems. To test this hypothesis, we measured the effects of simulated grazing on forage nitrogen, digestibility, and biomass. Simulated grazing increased average forage nitrogen and digestibility while decreasing forage biomass. These effects were associated with reduced individual growth rates, increased juvenile foraging time, and reduced juvenile vigilance. Results suggest that the negative effects of reduced vegetation biomass greatly outweighed positive treatment effects in this study. However, prairie dogs in the moderate intensity defoliation treatment showed some preference for “grazed” plots over “ungrazed” plots, and this preference increased with time. Our study lends support to the idea that habitat productivity and herbivore densities may mediate shifts between facilitative and competitive interactions between different-sized herbivores.     

The effect of sub-lethal increases in temperature on the growth and population trajectories of three scleractinian corals on the southern Great Barrier Reef

To date, coral death has been the most conspicuous outcome of warming tropical seas, but as temperatures stabilize at higher values, the consequences for the corals remaining will be mediated by their demographic responses to the sub-lethal effects of temperature. To gain insight into the nature of these responses, here I develop a model to test the effect of increased temperature on populations of three pocilloporid corals at One Tree Island, near the southern extreme of the Great Barrier Reef (GBR). Using Seriatopora hystrix, S. caliendrum and Pocillopora damicornis as study species, the effects of temperature on growth were determined empirically, and the dynamics of their populations determined under natural temperatures over a 6-month period between 1999 and 2000 [defined as the study year (SY)]. The two data sets were combined in a demographic test of the possibility that the thermal regime projected for the southern GBR in the next 55–83 years—warmer by 3°C than the study year (the SY+3 regime), which is equivalent to 1.4°C warmer than the recent warm year of 1998—would alter coral population trajectories through the effects on coral growth alone; the analyses first were completed by species, then by family after pooling among species. Laboratory experiments showed that growth rates (i.e., calcification) varied significantly among species and temperatures, and displayed curvilinear thermal responses with growth maxima at ∼27.1°C. Based on these temperature-growth responses, the SY+3 regime is projected to: (1) increase annualized growth rates of all taxa by 24–39%, and defer the timing of peak growth from the summer to the autumn and spring, (2) alter the intrinsic rate of population growth (λ) for S. hystrix (λ decreases 26%) and S. caliendrum (λ increases 5%), but not for P. damicornis, and (3) have a minor effect on λ (a 0.3% increase) for the Pocilloporidae, largely because λ varies more among species than it does between temperatures. Ten-year population projections suggest that the effects of a sub-lethal increase in temperature (i.e., the SY+3 regime) are relatively small compared to the interspecific differences in population dynamics, but nevertheless will alter the population size and increase the relative abundance of large colonies at the expense of smaller colonies for all three species, as well as the Pocilloporidae. These effects may play an important role in determining the nuances of coral population structure as seawater warms, and their significance may intensity if the coral species pool is depleted of thermally sensitive species by bleaching.     

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.     

Dynamics of an introduced population of mouflon Ovis aries on the sub‐Antarctic archipelago of Kerguelen

A commonly reported pattern in large herbivores is their propensity to irrupt and crash when colonizing new areas. However, the relative role of density‐dependence, climate, and cohort effects on demographic rates in accounting for the irruptive dynamics of large herbivores remains unclear. Using a 37‐yr time series of abundance in a mouflon Ovis aries population located on Haute Island, a sub‐Antarctic island of Kerguelen, 1) we investigated if irruptive dynamics occurred and 2) we quantified the relative effects of density and climate on mouflon population dynamics. Being released in a new environment, we expected mouflon to show rapid growth and marked over‐compensation. In support of this prediction, we found a two‐phase dynamics, the first phase being characterised by an irruptive pattern best described by the θ‐Caughley model. Parameter estimates were r_m=0.29±0.005(maximum growth rate), K=473±45 (carrying capacity) and S=2903±396 (surplus) mouflon. With a θ=3.18±0.69 our model also supported the hypothesis that density dependence is strongest at high density in large herbivores. The second phase was characterised by an unstable dynamics where growth rate was negatively affected by population abundance and winter precipitation. Climate, however, did not trigger population crashes and our model suggested that lagged density‐dependence and over‐grazing were the probable causes of mouflon irruptive dynamics. We compare our results with those of Soay sheep and discuss the possibility of a reversible alteration of the island carrying capacity after the initial over‐grazing period.     

Damage caused by low-density exotic herbivore populations: the impact of introduced European rabbits on marsupial herbivores and Allocasuarina and Bursaria seedling survival in Australian coastal shrubland

The impact of over-abundant exotic herbivores is well recognised, but their impact at low population densities is poorly understood. This study examined interactions between European rabbits and native herbivores, and their impact on seedling recruitment in coastal South Australia, 2 years after rabbit haemorrhagic disease (RHD) had reduced rabbit density to 4.48 rabbits ha⁻¹. Rabbit density was further reduced to 0.44 rabbits ha⁻¹ in replicated experimental treatments. Rabbit control reduced total grazing pressure by 39% despite compensatory grazing increases of >100% for both western grey kangaroos and common wombats. Rabbit control slowed the rate of grazing and mortality for planted drooping sheoak and sweet bursaria seedlings, but few survived for 12 months: 0 and 3% of sheoak, in untreated areas and rabbit control treatments, respectively, and 3 and 11% of bursaria, respectively. Planted sheoaks survived well if protected by rabbit-proof netting (60%). Within treatments, seedling grazing and survival rates were negatively correlated with rabbit density but kangaroo and wombat density had no measurable effect. We conclude that RHD may briefly have reduced rabbit densities enough to allow recruitment of bursaria but that sheoak require much lower rabbit densities than those provided by existing biological control agents. If left unaddressed, rabbit grazing could ultimately lead to the loss of sheoaks throughout most of their current range, irrespective of other attempts to conserve them. More generally, these data show how species-specific damage caused by low-density exotic herbivore populations may occur in the presence of more abundant but less-damaging native herbivores.     

The effect of domestic sheep on forage plants of wild reindeer; a landscape scale experiment

Domestic herbivores often compete with wildlife for limited resources, and on longer time-scales, grazing may also increase or decrease coverage of important food plants to wildlife affecting the threshold density for when competition can be expected. In Norway, there are growing concerns about the effect of releasing 2.1 million domestic sheep (Ovis aries) for summer grazing into areas hosting wild populations of alpine reindeer (Rangifer tarandus). We quantified the effect of sheep grazing (0, 25 and 80 sheep/km²) on the development in coverage and abundance of plants known to be important in the diet of reindeer during summer (vascular plants) and winter (lichens) within a fully replicated, landscape scale (2.7 km²) experiment. From 2001 to 2005, the sedge, Carex bigelowii, increased while the herb Solidago virgaurea decreased in frequency at high density of sheep relative to controls (both marginally non-significant). There was no marked development in Deschampsia flexuosa, Salix herbacea or Hierarcium alpinum that could be related to sheep grazing intensity. Lichen coverage and height both decreased at high density of sheep from 2002 to 2005. Effects of low grazing intensity were closer to controls than to high grazing intensity. Our study highlights that high sheep grazing intensity induce changes to the plant community that, at the same time, can improve the summer habitat and detriment the winter habitat to reindeer. Many wild reindeer populations are fragmented and may thus be limited by either summer or winter range. The effect of sheep grazing is predicted to vary accordingly. However, currently, we have limited ability to quantify how much this explicitly means in terms of increased or reduced carrying capacity for reindeer.     

Time-scale effects in the interaction between a large and a small herbivore

In the short term, grazing will mainly affect plant biomass and forage quality. However, grazing can affect plant species composition by accelerating or retarding succession at longer time-scales. Few studies concerning interactions among herbivores have taken the change in plant species composition into account. In a salt-marsh system, the long-term effects of exclusion of a large herbivore (cattle) on the abundance of a small herbivore (hare) were studied. Excluding cattle grazing for 30 years resulted in large changes in vegetation composition. In general, the cover of tall-growing species increased in the absence of cattle grazing. These long-term changes negatively affected hare grazing intensity. Hares preferentially fed on Festuca rubra and negatively selected tall growing plants, such as Elymus athericus, both in cattle-grazed and long-term ungrazed areas. However, the intensity of hare grazing was not related to the cover of F. rubra. The cover of tall-growing plants (E. athericus, Atriplex prostrata and Juncus maritimus) appeared to be the best predictor and hare grazing intensity decreased sharply with an increase of the cover of tall plants. When cover of tall plants did not increase, hare grazing intensity was not affected. The study shows that the time-scale of the experiment is of prime importance in studying interactions between herbivores. Species that do not seem to influence the abundance of one another or are competing for the same resources on a short time-scale might well be facilitating each other when looking at larger time-scales while taking plant species replacement into account.     

Effects of introduction and exclusion of large herbivores on small rodent communities

In this study we analysed the effects of large herbivores on smallrodent communities in different habitats using large herbivore exclosures. Westudied the effects of three year grazing introduction by red deer(Cervus elaphus L.) in previously ungrazed pine and oakwoodland and the exclusion of grazing by red deer, roe deer(Capreoluscapreolus L.) and mouflon (Ovis ammon musiminL.) in formerly, heavily grazed pine woodland and heathland. At eight exclosuresites within each habitat type, small rodents were captured with live trapsusing trapping grids. At each trapping grid, seed plots of beechnuts(Fagus sylvatica L.) and acorns (Quercusrobur L.) were placed to measure seed predation by rodents.Exclusion of grazing by large herbivores in formerly, heavily grazedhabitats had a significant effect on small rodent communities. Insideexclosureshigher densities of mainly wood mice (Apodemus sylvaticusL.) and field voles (Microtus agrestis L.) were captured.Introduction of grazing by red deer appeared to have no significant negativeeffects on small rodent communities. The seed predation intensity of beechnutsand acorns by small rodents was significantly higher in ungrazed situations,particularly in habitats that were excluded from grazing. The differencesbetween grazing introduction and exclusion effects on small rodent communitiescan be explained by differences in vegetation structure development. Therecovery of heavily browsed understory vegetation after large herbivore grazingexclusion proceeded faster than the understory degradation due to grazingintroduction. Small rodents depend on structural rich vegetations mainly forshelter. We conclude that large herbivores can have significant effects onvegetation dynamics not only via direct plant consumption but also throughindirect effects by reducing the habitat quality of small rodent habitats.     

内蒙古草原布氏田鼠与绵羊的营养生态位研究

放牧活动对小型啮齿动物种群数量的影响是草原生态系统研究中的热点问题,确定绵羊和小型啮齿动物的营养生态位关系对于草原生态系统的管理具有重要的作用。运用粪便显微分析法分析了内蒙古草原动物生态研究站大型野外围栏内布氏田鼠和绵羊在6—9月份的食谱组成,确定两种草食动物食性选择和营养生态位的变化,从食物利用途径揭示放牧活动对布氏田鼠种群数量的影响方式及二者之间的竞争与共存关系。研究结果表明：围栏中共有23种植物,其中糙隐子草、克氏针茅、羊草和刺藜是围栏中的优势植物;布氏田鼠共取食10种植物,绵羊共取食8种植物,禾本植物为两种动物的主要采食植物。布氏田鼠和绵羊的喜食植物种类都存在季节性变化;除7月外,两种动物的食物多样性与围栏植物多样性有显著的正相关关系,表明动物的食性选择受植物资源变化的影响。布氏田鼠和绵羊具有很高的营养生态位重叠度,除8月(0.691)外,6、7、9月份的营养生态位重叠度均高于0.9,表明二者之间存在激烈的食物竞争;放牧活动极显著地降低了禾本科植物的地上生物量,导致布氏田鼠的食物数量减少;放牧处理下布氏田鼠的喜食植物的物种数和营养生态位宽度(除9月)增加,导致布氏田鼠的食物质量...