The influence of soil depth on plant species response to grazing within a semi-arid savanna

Grassland patches within a semi-arid savanna were evaluated over 45-years for (1) local temporal dynamics of basal area for five dominant grass species within long-term heavily grazed and ungrazed treatments, (2) the influence of soil depth (resource availability) on vegetation dynamics, and (3) the applicability of community-level grazing response groups over fine-scale patterns of soil heterogeneity. Temporal patterns in species composition and basal area were dependent upon soil depth. In the heavy grazed treatment, Hilaria belangeri dominated deep soils while Erioneuron pilosum and Bouteloua trifida were restricted to shallow soils. In the ungrazed treatment, removal of grazing resulted in successional changes that were significantly different across soil depths. After 45 years without grazing, Eriochloa sericea was most abundant on deep soils while Bouteloua curtipendula was more abundant on intermediate and shallow soils. Community-level functional groups that are based on grazing were not appropriate when multiple pattern-driving variables were considered across multiple scales indicating that functional groups should only be applied to certain processes at specific scales. Within the ungrazed treatments, variable soil depths have resulted in a shifting mosaic in time and space where early- and late-successional species co-exist continuously but spatially separated within the community. In the heavily grazed treatment, species are somewhat spatially arranged by soil depths, but much of the inherent heterogeneity is eliminated and species composition is dominated by the three grazing-resistant short-grasses. Broad scale successional changes may appear linear and predictable while at finer scales, the same changes may be described as non-linear and dependent upon soil depth resulting in thresholds that are partially explained by weather patterns, seed bank limitations and competitive inhibitions.     

Grazing and an invasive grass confound spatial pattern of exotic and native grassland plant species richness

Previous work has shown exotic and native plant species richness are negatively correlated at fine spatial scales and positively correlated at broad spatial scales. Grazing and invasive plant species can influence plant species richness, but the effects of these disturbances across spatial scales remain untested. We collected species richness data for both native and exotic plants from five spatial scales (0.5–3000 m²) in a nested, modified Whittaker plot design from severely grazed and ungrazed North American tallgrass prairie. We also recorded the abundance of an abundant invasive grass, tall fescue (Schedonorus phoenix (Scop.) Holub), at the 0.5-m² scale. We used linear mixed-effect regression to test relationships between plant species richness, tall fescue abundance, and grazing history at five spatial scales. At no scale was exotic and native species richness linearly related, but exotic species richness at all scales was greater in grazed tracts than ungrazed tracts. Native species richness declined with increasing tall fescue abundance at all five spatial scales, but exotic species richness increased with tall fescue abundance at all but the broadest spatial scales. Severe grazing did not reduce native species richness at any spatial scale. We posit that invasion of tall fescue in this working landscape of originally native grassland plants modifies species richness-spatial scale relationships observed in less disturbed systems. Tall fescue invasion constitutes a unique biotic effect on plant species richness at broad spatial scales.     

Spatial heterogeneity and plant species richness at different spatial scales under rabbit grazing

Herbivores influence spatial heterogeneity in soil resources and vegetation in ecosystems. Despite increasing recognition that spatial heterogeneity can drive species richness at different spatial scales, few studies have quantified the effect of grazing on spatial heterogeneity and species richness simultaneously. Here we document both these variables in a rabbit-grazed grassland. We measured mean values and spatial patterns of grazing intensity, rabbit droppings, plant height, plant biomass, soil water content, ammonia and nitrate in sites grazed by rabbits and in matched, ungrazed exclosures in a grassland in southern England. Plant species richness was recorded at spatial scales ranging between 0.0001 and 150 m(2). Grazing reduced plant height and plant biomass but increased levels of ammonia and nitrate in the soil. Spatial statistics revealed that rabbit-grazed sites consisted of a mixture of heavily grazed patches with low vegetation and nutrient-rich soils (lawns) surrounded by patches of high vegetation with nutrient-poor soils (tussocks). The mean patch size (range) in the grazed controls was 2.1 +/- 0.3 m for vegetation height, 3.8 +/- 1.8 m for soil water content and 2.8 +/- 0.9 m for ammonia. This is in line with the patch sizes of grazing (2.4 +/- 0.5 m) and dropping deposition (3.7 +/- 0.6 m) by rabbits. In contrast, patchiness in the ungrazed exclosures had a larger patch size and was not present for all variables. Rabbit grazing increased plant species richness at all spatial scales. Species richness was negatively correlated with plant height, but positively correlated to the coefficient of variation of plant height at all plot sizes. Species richness in large plots (<25 m(2)) was also correlated to patch size. This study indicates that the abundance of strong competitors and the nutrient availability in the soil, as well as the heterogeneity and spatial pattern of these factors may influence species richness, but the importance of these factors can differ across spatial scales.     

Long-term vegetation dynamics mediated by herbivores,weather and fire in a Juniperus-Quercus savanna

Abstract. Long-term (45-yr) basal area dynamics of dominant graminoid species were analyzed across three grazing intensity treatments (heavily grazed, moderately grazed and ungrazed) at the Texas A&M University Agricultural Research Station on the Edwards Plateau, Texas. Grazing intensity was identified as the primary influence on long-term variations in species composition. Periodic weather events, including a severe drought (1951–1956), had little direct influence on composition dynamics. However, the drought interacted with grazing intensity in the heavily grazed treatment to exacerbate directional changes caused by grazing intensity. Species response to grazing was individualistic and noisy. Three response groups were identified. Taller, more productive mid-grasses were most abundant under moderate or no grazing. Short grasses were most abundant under heavy grazing. Intermediate species were most abundant under moderate grazing and opportunistic to weather patterns. Graminoid diversity increased with the removal or reduction of grazing intensity. The moderately and ungrazed treatments appeared most resistant to short-term weather fluctuations, while the heavily grazed treatment demonstrated significant resilience when grazing intensity was reduced after over 110 yr of overgrazing. Identification of a ‘climax’ state is difficult. Significant directional change, which took nearly 20 yr, appears to continue in the ungrazed treatment after 45 yr of succession. The observed, relatively linear patterns of perennial grass composition within the herbaceous patches of this savanna were generally explained by traditional Clementsian succession. However, when dynamics of the herbaceous community are combined with the woody component of this savanna, the frequency and intensity of fire becomes more important. Across the landscape, successional changes follow several pathways. When vegetation change is influenced by several factors, a multi-scale model is necessary to demonstrate interactions and feedbacks and accurately describe successional patterns. Absence of fires, with or without grazing, leads ultimately to a Juniperus/Quercus woodland with grazing intensity primarily influencing the fuel load and hence fire intensity.     

The impact of grazing management on Orthoptera abundance varies over the season in Mediterranean steppe-like grassland

As semi-natural grassland has a high level of biological diversity, understanding the effects of grazing and its variation over time is important in order to identify sustainable grazing practices. We measured temporal variation in Orthoptera abundance and spatial vegetation structure during seasonal grazing in an extensive sheep-farming system. We studied five grazed pasture areas (pre-grazing and post-grazing) and two adjacent ungrazed grasslands. We recorded the total abundance of Orthoptera and described the vegetation structure of 175 replicate plots (25 per pasture/grassland) during six field sampling sessions. We demonstrated that the impact of grazing on Orthoptera abundance is species-specific and greatly varies over the grazing season. The decrease of phytovolume is significant after 4–7 weeks of sheep grazing. Total Orthoptera abundance was higher in pre-grazed plots than in ungrazed plots, and higher in ungrazed plots than in post-grazed plots. These differences were particularly high during the peak of adult abundance. No difference in species richness was observed between grazing intensities. Total Orthoptera abundance positively correlated to phytovolume only when grazing pressure was high. However, the relationship between abundance and phytovolume differed between species. Extensive grazing by sheep tends to homogenize spatial vegetation structure and to temporarily reduce total Orthoptera abundance at pasture scale. However, rotational grazing allows spatial and temporal heterogeneity in vegetation structure to be maintained at farm scale, heterogeneity that is beneficial for Orthoptera. In contrast, absence of grazing has a negative impact on Orthoptera abundance as it favours the accumulation of litter, which is detrimental for a high proportion of xerothermophilic Orthoptera associated with bare ground and short vegetation.     

Broad‐scale patterns in plant diversity vary between land uses in a variegated temperate Australian agricultural landscape

Plant diversity is threatened in many agricultural landscapes. Our understanding of patterns of plant diversity in these landscapes is mainly based on small‐scale (<1000 m²) observations of species richness. However, such observations are insufficient for detecting the spatial heterogeneity of vegetation composition. In a case‐study farm on the North‐West Slopes of New South Wales, Australia, we observed species richness at four scales (quadrat, patch, land use and landscape) across five land uses (grazed and ungrazed woodlands, native pastures, roadsides and crops). We applied two landscape ecological models to assess the contribution of these land uses to landscape species richness: (i) additive partitioning of diversity at multiple spatial scales, and (ii) a measure of habitat specificity – the effective number of species that a patch contributes to landscape species richness. Native pastures had less variation between patches than grazed and ungrazed woodlands, and hence were less species‐rich at the landscape scale, despite having similar richness to woodlands at the quadrat and patch scale. Habitat specificity was significantly higher for ungrazed woodland patches than all other land uses. Our results showed that in this landscape, ungrazed woodland patches had a higher contribution than the grazed land uses to landscape species richness. These results have implications for the conservation management of this landscape, and highlighted the need for greater consensus on the influence of different land uses on landscape patterns of plant diversity.     

The effects of grazing on local colonisation and extinction during early succession

Estimates of colonisation of plant species were made at three spatial scales in an old-field on limestone subject to five experimental sheep grazing regimes. Local extinctions within grazing treatments were estimated in 1-m² permanent quadrats. These data were used to assess the effects of grazing treatment and spatial scale on the process of species change over a period of six years. Colonisation of the lOha field was virtually a random draw of plant species from adj acent vegetation, irrespective of plant life-history traits including dispersal strategy. The effects of grazing on colonisation increased at smaller spatial scales. Colonisation rates changed little during the study on the 10 ha scale, but declined steeply with time at smaller scales. Colonisation rates of short-lived species declined more than those of perennials, and short-lived species were subject to erratic episodes of extinction which did not affect perennials. Short periods of grazing enhanced colonisation rates of all species, but extinction rates were the same as in ungrazed controls. Grazing for longer periods further enhanced colonisation rates, but also increased extinction rates. This produced diversity patterns consistent with a ‘hump-backed model’, except that no grazing treatment was heavy enough to decrease diversity. Ungrazed controls had low species diversity, but areas grazed for longer periods were no more diverse after six years than those grazed for short periods.     

Livestock exclusion increases the spatial heterogeneity of vegetation in Colorado shortgrass steppe

Abstract. Spatial heterogeneity, an important characteristic in semi‐arid grassland vegetation, may be altered through grazing by large herbivores. We used Moran's I, a measure of autocorrelation, to test the effect of livestock grazing on the fine scale spatial heterogeneity of dominant plant species in the shortgrass steppe of northeastern Colorado. Autocorrelation in ungrazed plots was significantly higher than in grazed plots for the cover of the dominant species Bouteloua gracilis, litter cover and density of other bunchgrasses. No species had higher autocorrelation in grazed compared to ungrazed sites. B. gracilis cover was significantly auto‐correlated in seven of eight 60‐yr ungrazed exclosures, four of six 8‐yr exclosures, and only three of eight grazed sites. Autocorrelograms showed that B. gracilis cover in ungrazed sites was frequently and positively spatially correlated at lag distances less than 5 m. B. gracilis cover was rarely autocorrelated at any sampled lag distance in grazed sites. The greater spatial heterogeneity in ungrazed sites appeared linked to patches characterized by uniformly low cover of B. gracilis and high cover of C₃ grasses. This interpretation was supported by simple simulations that modified data from grazed sites by reducing the cover of B. gracilis in patches of ca. 8 m diameter and produced patterns quite similar to those observed in ungrazed sites. In the one exclosure where we intensively sampled soil texture, autocorrelation coefficients for sand content and B. gracilis cover were similar at lag distances up to 12 m. We suggest that the negative effect of sand content on B. gracilis generates spatial heterogeneity, but only in the absence of grazing. An additional source of heterogeneity in ungrazed sites may be the negative interaction between livestock exclusion and B. gracilis recovery following patchy disturbance.     

A hierarchical model for analysing the stability of vegetation patterns created by grazing in temperate pastures

Questions: Does vegetation structure display any stability over the grazing season and in two successive years, and is there any correlation between the stability of these spatial patterns and local sward composition? Location: An upland grassland in the French Massif Central. Method: The mosaic of short and tall vegetation stands considered as grazed and ungrazed patches respectively is modeled as the realization of a Boolean process. This method does not require any arbitrarily set sward‐height thresholds to discriminate between grazed and ungrazed areas, or the use of additional variables such as defoliation indexes. The model was validated by comparing empirical and simulated sward‐height distributions and semi‐variograms. Results: The model discriminated between grazed and ungrazed patches at both a fine (1 m²) and a larger (500 m²) scale. Selective grazing on legumes and forbs and avoidance of reproductive grass could partly explain the stability of fine‐scale grazing patterns in lightly grazed plots. In these plots, the model revealed an inter‐annual stability of large‐scale grazing patterns at the time peak biomass occurred. At the end of the grazing season, lightly grazed plots showed fluctuating patch boundaries while heavily grazed plots showed a certain degree of patch stability. Conclusion: The model presented here reveals that selective grazing at the bite scale could lead to the creation of relatively stable patches within the pasture. Locally maintaining short cover heights would result in divergent within‐plot vegetation dynamics, and thus favor the functional diversity of vegetation.     

From plant neighbourhood to landscape scales: how grazing modifies native and exotic plant species richness in grassland

The interactive effect of grazing and soil resources on plant species richness and coexistence has been predicted to vary across spatial scales. When resources are not limiting, grazing should reduce competitive effects and increase colonisation and richness at fine scales. However, at broad scales richness is predicted to decline due to loss of grazing intolerant species. We examined these hypotheses in grasslands of southern Australia that varied in resources and ungulate grazing intensity since farming commenced 170 years ago. Fine-scale species richness was slightly greater in more intensively grazed upper slope sites with high nutrients but low water supply compared to those that were moderately grazed, largely due to a greater abundance of exotic species. At broader scales, exotic species richness declined with increasing grazing intensity whether nutrients or water supply were low or high. Native species richness declined at all scales in response to increasing grazing intensity and greater resource supply. Grazing also reduced fine-scale heterogeneity in native species richness and although exotics were also characterised by greater heterogeneity at fine scales, grazing effects varied across scales. In these grasslands patterns of plant species richness did not match predictions at all scales and this is likely to be due to differing responses of native and exotic species and their relative abundance in the regional species pool. Over the past 170 years intolerant native species have been eliminated from areas that are continually and heavily grazed, whereas transient, light grazing increases richness of both exotics and natives. The results support the observation that the processes and scales at which they operate differ between coevolved ungulate—grassland systems and those in transition due to recent invasion of herbivores and associated plant species.     

Impact of grazing on soil carbon and microbial biomass in typical steppe and desert steppe of Inner Mongolia

The potential of grazing lands to sequester carbon must be understood to develop effective soil conservation measures and sustain livestock production. Our objective was to evaluate the effects of grazing on soil organic carbon (SOC), total nitrogen (TN), microbial biomass carbon (MBC) in Typical steppe and Desert steppe ecosystems, which are both important grassland resources for animal grazing and ecological conservation in China, and to derive region-specific soil C changes associated with different stocking rates (ungrazed, UG; lightly grazed, LG; moderately grazed, MG; heavily grazed, HG). This study substantiated that significant higher SOC, TN and MBC appeared with the treatment of LG in typical steppe. From 2004 to 2010, grazing treatments increased soil carbon storage in desert steppe, which was partly due to the grazing history. The higher MBC concentration and MBC/SOC suggest a great potential for carbon sequestration in the desert steppe ecosystem. The greater MBC in desert steppe than typical steppe was mainly the result of higher precipitation and temperature, instead of soil substrate. The change of MBC and the strong positive relationships between MBC and SOC indicated that MBC in the soil was a sensitive index to indicate the dynamics of soil organic carbon in both steppes in Inner Mongolia of China.     

Birds bias offspring sex ratio in response to livestock grazing

Livestock grazing, which has a large influence on habitat structure, is associated with the widespread decline of various bird species across the world, yet there are few experimental studies that investigate how grazing pressure influences avian reproduction. We manipulated grazing pressure using a replicated field experiment, and found that the offspring sex ratio of a common upland passerine, the meadow pipit Anthus pratensis, varied significantly between grazing treatments. The proportion of sons was lowest in the ungrazed and intensively grazed treatments and highest in treatments grazed at low intensity (by sheep, or a mixture of sheep and cattle). This response was not related to maternal body condition. These results demonstrate the sensitivity of avian reproductive biology to variation in local conditions, and support growing evidence that too much grazing, or the complete removal of livestock from upland areas, is detrimental for common breeding birds.     

Effects of grazing on soil seed bank dynamics: An approach with functional groups

Abstract. The relationship between intensity and timing of cattle grazing on changes in the size and composition of the soil seed bank were investigated in a 3‐yr study in a Mediterranean grassland in northeastern Israel. Treatments included manipulations of stocking rates and of grazing regimes, in a factorial design. The retrieved soil seed bank community was rich in species, with 133 species accounting for 80% of the 166 species recorded at the site. Within the seed bank, 89% of the species were annuals. Seed bank dynamics was analysed in terms of plant functional groups and germination strategies. In terms of total seed bank density and including all functional groups, 42% of the seeds present in the soil did not germinate under watering conditions. The dormancy level differed greatly among functional groups. The seed bank of annual legumes, crucifers, annual thistles and annual forbs had a large fraction of non‐germinated seeds and characterized areas grazed early in the growing season under high and very high grazing intensity. These functional groups were considered to have a higher potential for persistent seed banks production. In contrast, short and tall annual grasses and tall perennial grasses, that were dominant in ungrazed or moderately grazed paddocks, generally had seed banks with a very small fraction of non‐germinated seeds. Seed bank densities varied widely between grazing treatments and years. Under continuous grazing, heavy grazing pressure reduced seed bank densities of grasses and crucifers in comparison to moderate grazing. The greatest reduction on the seed bank densities resulted from heavy grazing concentrated during the seed‐set stages.     

Effects of algivorous minnows (Campostoma) on spatial and temporal heterogeneity of stream periphyton

The effect of grazing minnows (Campostoma) on spatial and temporal heterogeneity of the vertical height of attached filamentous algae (Spirogyra and Rhizoclonium) was measured in natural and artificial streams. Measurements were made at 1-m intervals across natural-stream pools during April–May, and at 0.3-m intervals longitudinally in smaller artificial streams during February–May. Spatial heterogeneity was calculated weekly, as the mean standardized difference in algal height (spatial MDH) between adjacent fixed points. Temporal MDH was calculated as the mean standardized difference in algal height at fixed points between adjacent weeks. Reduction in spatial and temporal MDH, detected only in artificial streams, suggested that grazing by Campostoma promoted and maintained more uniform algal height in contrast to ungrazed algae. Heterogeneity of algal export was greater for ungrazed algae, and decreased over time after sloughing, but that for grazed algae was lower and increased after sloughing. The contrasting experimental systems suggested that Campostoma can reduce spatial and temporal heterogeneity of algae, but that additional variation in depth, substratum characteristics, or presence of other biota may modify effects in natural streams. Received: 30 May 1995 / Accepted: 30 June 1997     

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.     

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.     

The effect of grazing on granivory patterns in the temperate Monte Desert, Argentina

Structural variations result in a marked heterogeneity of processes in arid environments. Disturbances have noticeable impacts on ecosystems, and a solid way of assessing their effect is to analyze how ecological processes operate. In deserts, granivory has been considered as a keystone process. The objective of our study was to determine the effect of domestic grazing on seed removal rates by birds, ants and rodents as an indirect measure of the process of granivory. This study was carried out in grazed and ungrazed habitats in the central Monte desert ecoregion in Argentina, during February and July 2002. We analyzed three major habitats: mesquite forest, creosotebush shrubland, and sand dunes under two different treatments (grazed, ungrazed). We found significant effects of grazing and non-grazing treatments on total seed removal rates, among granivorous taxa, as well as interactions with season and habitats. Ants and birds were the taxa that differed most under grazing pressure whereas seed removal by rodents showed no significant responses towards grazing. In conclusion, grazing affects plant structure and the assemblage of granivorous taxa, which in turn translates to the granivory process, adding another source of variation to the seed removal patterns reported for the temperate Monte desert.     

Importance of grazing and soil acidity for plant community composition and trait characterisation in coastal dune grasslands

Question: Does grazing by large herbivores affect species composition or community‐wide variation in plant functional traits? Location: Dune grasslands at the Belgian coast. Methods: Plant cover and soil data were collected in 146 plots that were randomly selected at 26 grazed and ungrazed grassland sites. Plant community composition was assessed by Detrended Correspondence Analysis and mean values of plant trait categories were calculated across the plots. Results: Differentiation of plant composition and community‐wide plant trait characteristics was largely determined by grazing, soil acidity and their interaction. In ungrazed situations, a clear floristic distinction appears between acidic (non‐calcareous) and alkaline (calcareous) grasslands. In grazed situations, these floristic differences largely disappeared, indicating that grazing results in a decrease of natural variation in species composition. At higher soil pH, a larger difference in plant community composition and community‐wide plant traits was observed between grazed and ungrazed plots. In ungrazed situations, shifts in plant functional traits along the acidity gradient were observed. Conclusions: Grazing is responsible for shifts in plant community composition, and hence a decrease in plant diversity among grasslands at opposing acidity conditions in coastal dune grasslands. Therefore, care should be taken when introducing grazing as a system approach for nature conservation in dune grasslands as it may eliminate part of the natural variation in plant diversity along existing abiotic gradients.     

Impact of reindeer grazing on ground-dwelling Carabidae and Curculionidae assemblages in Lapland

Reindeer Rangifer tarandus L. grazing shapes forest vegetation, microclimate, and soil respiration in Lapland, especially due to grazing on lichens (Cladina). We studied how these changes and their magnitude affect ground‐dwelling species of beetle families Carabidae (predators) and Curculionidae (herbivores), by using pitfall traps to collect invertebrates from pairs of grazed and ungrazed study plots over a wide range of site types. Changes in abundance, composition, richness and diversity of beetle assemblage were tested in relation to magnitude of the impacts on vegetation. The species compositions of Carabidae and Curculionidae differed between grazed and ungrazed plots in all sites. The relative difference between grazed and ungrazed plots in the number of individuals increased linearly with the impact of reindeer on vegetation cover. Carabid beetles, as a family, were more common in grazed plots in all sites. Curculionid beetles were more common in ungrazed plots in the birch dominated sites. This difference was mainly due to the species that feeds on deciduous leaves. In the pine dominated sites with high Cladina cover and more changes in ground vegetation, the number of curculionids feeding on conifers was higher in grazed plots. Species richness and diversity (H’) of both families were higher in grazed plots. Of the total 27 species, 11 were found only in grazed plots, while not a single species was found only in ungrazed plots. The relative difference between plots in diversity and evennes (H’/H'max) had humped response to the difference in Cladina cover. The diversity values were greater in grazed plots at the intermediate levels of grazing impact, and only in sites with very low or extremely high Cladina cover difference was the diversity higher in ungrazed plots. The response of beetle diversity resembled the hypotheses suggested for the relationship between grazing and vegetation diversity: greatest positive effect at intermediate grazing intensity and negative effects at unproductive sites.