Long-term Exclosure Research Articles

Plant community dynamics in shortgrass steppe with grazing relaxation and imposition by large and small herbivores

Semiarid rangelands often respond slowly to rest/relaxation of grazing pressure by large herbivores, and the effects of grazing are most often inferred from this direction of study because the imposition of grazing onto previous ungrazed/lightly grazed areas occurred prior to the age of scientific studies. These rangelands host a diversity of small and large herbivores, but grazing studies most often concern effects of the large generalists. Here, the effects of herbivore body size on plant species richness and dominant species, and imposition and relaxation of grazing by large herbivores were studied by opening half of exclosures established in 1939 and building new exclosures to large herbivores, and to small-plus-large herbivores. Plant richness using sensitive species-area sampling was studied in a dry and a wet year, about 62 years after initiating the long-term experiment and about 6-10 years after initiating the altered designs. Convergence of the newly opened to large herbivore grazing treatment to the long-term grazed treatment occurred within 10 years, but convergence of the newly excluded to large herbivore treatment to the long-term excluded treatment had only partly occurred. This indicated that recovery from grazing is slow relative to imposition of grazing by large herbivores, but effects of the additional exclusion of small-plus-large herbivores occurred relatively rapidly. These results were mirrored by trajectories of convergence of the dominant species, and this is discussed with respect to implications for state-and-transition models. Short-term exclusion of small-plus-large herbivores resulted in greater richness than even long-term exclusion of only large herbivores, even though quantities consumed by small herbivores are much less than by large. Grazing effects on plant richness were large in the wet year, but the very dry year suppressed richness on all treatments. When sampling effort and area are the same, the numbers and attributes of species unique to a treatment are indicators of rareness of the richness and traits selected for by the treatment. More unique species were sampled in the small-plus-large herbivore exclosures when comparing body size, and the long-term large herbivore exclosures when comparing time of exclosure. Unique species encountered during sampling the ungrazed treatments were generally forbs, exotic and/or weedy invasive species, and often tall, annual species.

Capability Investigation of Carbon Sequestration in Two Species (Artemisia sieberi Besser and Stipabarbata Desf) Under Different Treatments of Vegetation Management (Saveh, Iran)

The rangelands, as one of the largest dynamic biomes in the world, have very capabilities. Regulation of greenhouse gases in the Earth's atmosphere, particularly carbon dioxide as the main greenhouse gases, is one of these cases. The attention to rangeland, as cheep and reachable resources to sequestrate the carbon dioxide, increases after the Industrial Revolution. Rangelands comprise the large parts of Iran as a steppic area. Rudshur (Saveh), as area index of steppic area, was selected under three sites including long-term exclosure, medium-term exclosure, and grazable area due to the capability of carbon dioxide's sequestration of dominated species. Canopy cover's percentage of two dominated species (Artemisia sieberi Besser and Stipa barbata Desf.) was determined via establishing of random 1 square meter plot. The sampling of above and below ground biomass style was obtained by complete random. After determination of ash percentage in the laboratory; conversion ratio of plant biomass to organic carbon was calculated by ignition method. Results of the paired t-test showed that the amount of carbon sequestration in above ground and underground biomass of Artemisia sieberi Besser and Stipa barbata Desf. is different in three regions. It, of course, has not any difference between under and surface ground's biomass of Artemisia sieberi Besser in long-term exclosure. The independent t-test results indicate differences between underground biomass corresponding each other in the studied sites. Carbon sequestration in the Stipa barbata Desf. was totally more than Artemisia sieberi Besser. Altogether, the average sequestration of the long- term exclosure was 5.842gr/m², the medium-term exclosure was 4.115gr/m², and grazable area was 5.975gr/m² so that there is not valuable statistical difference in terms of total amount of carbon sequestration to three sites.

THE EFFECTS OF EXCLOSURE ON PLANTS IN THE SEMI-ARID RANGELAND OF NORTH KHORASAN PROVINCE, IRAN

Abstract Over the last fifty years, most of the semi-arid rangeland in Iran has been converted to cropland without an equivalent reduction in grazing animals. This shift has led to heavy grazing pressure on rangeland vegetation. The present study has been conducted in Sisab Research Station in the North Khorasan Province to evaluate effects of long-term grazing exclosures on biodiversity of range plants. The study was initiated in May 2008 using three transects within the exclosure and another three transects near exclosure, each with 100m length in the vicinity of exclosure has been established to determine differences in plant composition between areas that have not been grazed in 22 years with neighboring grazed plant communities. The sampling method was randomized systematic one comprised of 10 plots, each of 1m2 in area. A total of 53 plant species were identified in the study area with the ungrazed plots containing 18 plants more than the grazed plots. The major species were Festuca ovina, Centurea depressa, Stachys turkamanica, Stipa barbata, Astragalus sp and Phelomis cancellata. Grazing impacts on forbs were more pronounced than for grasses and shrubs. Based on Jaccard’s index, there was only a 45% similarity of plant species between the two treatments. Our study led to four generalizations about the current grazing regime and long-term exclosures in the semi-arid rangeland around the study area: (1) exclosures will increase species richness, (2) heavy grazing may have removed some plant species, (3) complete protection from grazing for a prolonged period of time after a long history of grazing disturbance may not lead to an increase in desirable plant species with a concomitant improvement in range condition, and (4) research needs to be conducted to determine how these rangelands can be improved.

Do mammalian herbivores influence invertebrate communities via changes in the vegetation? Results from a preliminary survey in Kruger National Park, South Africa

We investigated the indirect influence of mammalian herbivores on invertebrates, by utilising long-term mammalian herbivore exclosures in Kruger National Park. The exclosures span three distinct habitat types (crest, footslope and riparian) on a catena. By performing invertebrate collections in the exclosures and in a control area we were able to assess the indirect effect of mammalian herbivory on the invertebrate community and if this influence varied across habitat types. Our results indicate that large mammalian herbivores (notably elephants) had significant negative effects on total invertebrate abundance, while medium-sized mammalian herbivores affected the abundance and richness of beetles and grasshoppers negatively. Habitat type affected the invertebrates; spider abundance and richness peaked on the footslope, while beetles were the most abundant and taxon-rich in the riparian zone. Hence, our results suggest that indirect effects of mammalian herbivores on the invertebrate community may be significant, and in most cases negative, but also that the effects of mammalian herbivores vary across invertebrate groups. Thus, to better understand the broad-scale implications of changed mammalian herbivore pressures for the functioning of savanna systems, it may be important to take effects on invertebrate communities into account.

Recovery of plant species composition and ecosystem function after cessation of grazing in a Mediterranean grassland

Short- and long-term changes in species composition, plant biomass production, and litter decomposition after cessation of grazing were examined in a Mediterranean grassland with high dominance of annual species and strong seasonality in biomass production. Short-term changes were assessed during three consecutive years in plots previously exposed to different grazing pressures and compared to plots in long-term (30–40 years) exclosures. Short-term cessation of grazing led in the short-term to an increase in relative biomass of annual crucifers and tall annual and perennial grasses, while biomass of annual legumes, annual thistles and short annual grasses decreased. Consequently, similarity increased between vegetation recently excluded from grazing and vegetation in long-term protected plots. Our research showed that in systems with high dominance of grasses and annual species, the rapid changes in plant species composition that occur after grazing cessation were associated with a fast recovery of the potential for biomass production to levels found in long-term protected plots, while litter decomposition rate did not change even after long-term cessation of grazing. Moreover, previous history of grazing did not affect plant litter decomposition, despite higher litter quality in grazed treatments. This study provides new insights about the processes involved in the diverse responses of ecosystem functions resulting from shifts in species composition associated with grazing cessation and land use change in Mediterranean grasslands.

Rainfall variability may modify the effects of long-term exclosure on vegetation in Mandalgobi, Mongolia

Starting in 2005, we examined differences in vegetation for three consecutive years across an airport fence that separated heavily grazed areas from areas in which grazing had been excluded for 24 years in Mandalgobi, Mongolia. We performed repeated-measures analysis separately on two community types (dominated by Allium polyrrhizum and Achnatherum splendens, respectively) to compare the effects of fencing and year on the cover of different plant functional types. There was a significant fence × year interaction for grass cover in the Allium type (but not the Achnatherum type), due to greater cover of grasses inside the fence only when rainfall was sufficient during the growing season. The effect of grazing exclusion on perennial forb cover was confounded by a significant fence × year interaction in both types. In 2007, perennial forbs were found outside the fence, but had almost disappeared inside the fence, resulting in this interaction. Annual forbs only had much greater cover values inside the fence than outside in 2006, also resulting in a significant fence × year interaction in both community types. This study thus suggests that the high rainfall variability in arid and semi-arid rangelands may modify the effects of long-term exclosure on vegetation.

Above-ground herbivory causes rapid and sustained changes in mycorrhizal colonization of grasses

Arbuscular mycorrhizal fungi (AMF) play a vital role in ecosystem functioning. In most grasslands, herbivory by both vertebrate and invertebrate herbivores is common and thus in order to assess herbivore effects on multitrophic-level interactions both should be considered. This study investigated the effects of grazing by rabbit and insect herbivores on root-colonization of grasses by AMF in two lowland grasslands in southern England, UK. A long-term exclosure site was used to provide a temporal assessment in order to elucidate whether any short-term responses to herbivore removal were sustained. Root samples from three grass species at each site were analysed in terms of total mycorrhizal colonization and proportional colonization by individual mycorrhizal structures. Colonization levels were up to 1.6 times greater under moderate levels of rabbit grazing (with summer maxima of 25% and winter minima of 11%) than in intensely grazed swards or fenced plots at both sites. The change was fast (within 8 weeks), consistent throughout the sampled field plots, and temporally sustainable over a 19-year period. There was no significant effect of insect herbivory on total colonization but proportional colonization by different AM structures was affected on some sample dates where vertebrate herbivores had been removed, indicating a slight effect on fungal structure allocation. The results suggest that the type of herbivore and perhaps more importantly the intensity of grazing are key determinants of below-ground effects upon mycorrhizal-host plant symbiosis. The data suggest that the extent of mycorrhizal colonization within grass host plants is strongly influenced by C assimilation and allocation.

The Effects of Long-Term Grazing Exclosures on Range Plants in the Central Anatolian Region of Turkey

Over the last fifty years, almost half of the steppe rangeland in the Central Anatolian Region of Turkey (CAR) has been converted to cropland without an equivalent reduction in grazing animals. This shift has led to heavy grazing pressure on rangeland vegetation. A study was initiated in June 2003 using 6 multiscale Modified-Whittaker plots to determine differences in plant composition between areas that have not been grazed in 27 years with neighboring grazed plant communities. A total of 113 plant species were identified in the study area with the ungrazed plots containing 32 plants more than the grazed plots. The major species were Astragalus acicularis, Bromus tomentellus, Festuca valesiaca, Genista albida, Globularia orientalis, Poa bulbosa, and Thymus spyleus ssp rosulans. Grazing impacts on forbs were more pronounced than for grasses and shrubs. Based on Jaccard's index, there was only a 37% similarity of plant species between the two treatments. Our study led to four generalizations about the current grazing regime and long-term exclosures in the steppe rangeland around the study area: (1) exclosures will increase species richness, (2) heavy grazing may have removed some plant species, (3) complete protection from grazing for a prolonged period of time after a long history of grazing disturbance may not lead to an increase in desirable plant species with a concomitant improvement in range condition, and (4) research needs to be conducted to determine how these rangelands can be improved.

INFLUENCE OF GRAZING BY LARGE MAMMALS ON THE SPIDER COMMUNITY OF A KENYAN SAVANNA BIOME

Abstract Pitfall trap and sweep net samples were taken over a period of fifteen months (2002– 2003) in the Kenya Long-term Exclosure Experiment (KLEE), in which the presence of domestic and wild herbivores have been independently manipulated since 1995. ANOVA and ANCOVA showed that the exclosure treatments significantly affected plant cover, with the presence of cattle significantly reducing the relative vegetation cover and spider diversity. Herbivory by indigenous mega- and meso–herbivores did not have a significant influence on the diversity of the spider fauna, but abundance of three dominant species (Cyclosa insulana Costa (Araneidae), Argiope trifasciata Forskal (Araneidae) and Runcinia flavida Simon (Thomisidae)) decreased in cattle-grazed plots. In contrast, Aelurillus sp. became more prevalent where cattle have been grazing. Multivariate analyses revealed that the spider community responded to grazing pressure by aggregating into three groups that reflected control, cattle grazing and non-cattle ...

Grazing History Affects Willow Communities in a Montane Riparian Ecosystem

This study was conducted to compare data from 12 grazed and ungrazed areas and to examine the impacts of grazing treatments on a montane willow community during an 11-year period. Data were collected on willow canopy cover, species diversity, height, and stem density in a montane riparian ecosystem between 1988 and 1999 from 4 grazing treatments: long-term grazing (since the early 1900s), long-term grazing exclusion (exclosures built in the 1950s), recent grazing (sections of exclosures opened in 1988), and recent grazing exclusion (exclosures built in 1988). Willow canopy cover increased significantly for all treatments through time, with the recent grazing exclusion treatment becoming similar to that of the long-term exclusion treatment within 5 years. Species diversity was greatest in the long-term grazed treatment. Willow height averaged over treatments increased from 1988 to 1997 (P = 0.0001), but did not increase significantly after that. Height in the long-term exclosure averaged over time from 1988 to 1997 was 1.5 times greater than in the long-term grazing treatment. Stem density of willows was significantly greater in the recent exclosure than in the long-term exclosure (P = 0.008, 180%) and recent grazing treatments (P = 0.02, 120%). Recent grazing exclusion resulted in the greatest increase in canopy cover, height growth, and stem density during the 11 years of study, indicating that these variables respond positively to removal of livestock grazing. Results suggest that continued long-term grazing exclusion may lead to a closed canopy, lower willow species diversity, reduction in new stem height growth, and reduced stem recruitment. Information on the dynamics of willow growth under different grazing treatments should help resource managers determine appropriate livestock utilization levels in similar riparian areas, and develop management plans for these important ecosystems.

Grazing History Affects Willow Communities in a Montane Riparian Ecosystem

This study was conducted to compare data from 12 grazed and ungrazed areas and to examine the impacts of grazing treatments on a montane willow community during an 11-year period. Data were collected on willow canopy cover, species diversity, height, and stem density in a montane riparian ecosystem between 1988 and 1999 from 4 grazing treatments: long-term grazing (since the early 1900s), long-term grazing exclusion (exclosures built in the 1950s), recent grazing (sections of exclosures opened in 1988), and recent grazing exclusion (exclosures built in 1988). Willow canopy cover increased significantly for all treatments through time, with the recent grazing exclusion treatment becoming similar to that of the long-term exclusion treatment within 5 years. Species diversity was greatest in the long-term grazed treatment. Willow height averaged over treatments increased from 1988 to 1997 (P = 0.0001), but did not increase significantly after that. Height in the long-term exclosure averaged over time from 1988 to 1997 was 1.5 times greater than in the long-term grazing treatment. Stem density of willows was significantly greater in the recent exclosure than in the long-term exclosure (P = 0.008, 180%) and recent grazing treatments (P = 0.02, 120%). Recent grazing exclusion resulted in the greatest increase in canopy cover, height growth, and stem density during the 11 years of study, indicating that these variables respond positively to removal of livestock grazing. Results suggest that continued long-term grazing exclusion may lead to a closed canopy, lower willow species diversity, reduction in new stem height growth, and reduced stem recruitment. Information on the dynamics of willow growth under different grazing treatments should help resource managers determine appropriate livestock utilization levels in similar riparian areas, and develop management plans for these important ecosystems.

Growth of Prosopis caldenia Burk: seedlings in central semi-arid rangelands of Argentina

Prosopis caldenia Burk. is one of the woody species increasing in abundance due to poor grazing management in the semi-arid regions of central Argentina. The objective of this study was to evaluate the effect of herb cover, cattle dung, soil compaction, and water supply on P. caldenia seedling growth on sites with different grazing history: a site exposed to a long-term (25 or more years) continuous grazing by cattle (grazed experimental site), and a site long-term (around 20 years) exclosure to domestic livestock (ungrazed experimental site). In both experimental sites, regardless of the study period, the highest height and main root length of P. caldenia seedlings were recorded in plots with the herb cover removed, soil disturbed, cattle dung presence, and water supply. Above- and below-ground biomass of P. caldenia seedlings showed a similar response. Direct and indirect effects of livestock grazing favoured P. caldenia seedlings growth, which might increase their establishment probabilities in sites under a chronic heavy grazing. Consequently, this might alter the grass-woody plant balance in the Caldenal causing a drastic reduction of the domestic livestock production in the region.

Factors related to establishment of Prosopis caldenia Burk. seedlings in central rangelands of Argentina

Prosopis caldenia Burk. is one of the woody species that is increasing in abundance due to poor grazing management in the semi-arid phytogeographic region of central Argentina, commonly known as the Caldenal. The objective of this study was to evaluate the effect of herbaceous cover, cattle dung, soil disturbance, and water supply on emergence and survival of P. caldenia seedlings on sites with different grazing histories: (i) a site exposed to long-term continuous grazing by cattle (grazed site), and (ii) a long-term exclosure to domestic livestock (ungrazed site). Removal of grass cover, addition of cattle dung, and water supply enhanced seedling emergence and survival, especially in the grazed site. Results suggest that factors (direct and indirect) associated with prolonged grazing history markedly affect P. caldenia establishment. This in turn alters the grass-woody plant balance, which might reduce the potential capacity of livestock production in the Caldenal.

Competition and compensation among cattle, zebras, and elephants in a semi-arid savanna in Laikipia, Kenya

Competition and compatibility between livestock and wildlife in Africa has been a point of considerable speculation, with implications for conservation. However, controlled replicated experiments are lacking. Here we report on the results of a long-term exclosure experiment in Laikipia, Kenya, in which different guilds of large mammalian herbivores have been independently manipulated since 1995. In plots from which cattle were excluded, the density of zebra dung in 2000 was on average 46% greater than in control plots. This was due to differential zebra use, and not to differential rates of dung removal (by dung beetles or other factors). Vegetation data indicate that cattle fully compensate for the absence of wildlife; all plots accessible to cattle had similarly low grass cover. However, wildlife do not fully compensate for the absence of cattle; plots with only wildlife had more grass cover than plots accessible to cattle. Zebra dung density was strongly correlated with total grass cover, suggesting that zebras are effectively tracking resource abundance. There is also evidence of pair-wise competition between cattle and elephants, and between elephants and zebras. The strong competition between cattle and zebras appears to be mitigated by the presence of elephants. A significant cattle x elephant interaction on the abundance of zebra dung indicates that elephants reduce the negative effects of cattle on zebras. In the presence of cattle, elephants facilitate the abundance of zebra, apparently by suppressing resource extraction (bite rates) by cattle. The precise mechanism for this indirect facilitation is not clear, but it may be related to the demonstrated reduction in forb cover associated with elephant presence.

Defoliation Effects on Reproductive Biomass: Importance of Scale and Timing

Community-level (per unit area) and individual tiller reproductive biomass inside and outside of long-term exclosures on the northern winter range of Yellowstone National Park, USA were compared. Grazed areas had twice the number of reproductive tillers m-2 (186 compared to 88 tillers m-2), and greater total reproductive biomass m-2 than ungrazed plots (13 compared to 7 g m-2). In contrast, seed number tiller-1 was greater for grasses in exclosures. Because of these offsetting responses, seed production (no. m-2) was unaffected by herbivores. On an area basis, grazed grasses allocated proportionally more biomass to reproduction (reproductive biomass/aboveground biomass) than ungrazed grasses. We propose that altered plant demography and morphology following defoliation explain how grazers might increase the allocation of biomass to reproduction in Yellowstone grasslands. To understand these results in light of ecological and agronomic studies, we reviewed literature from 118 sources that reported the effects of defoliation on the production of reproductive biomass. The review suggested that the results of herbivory or defoliation on plant reproductive biomass depended on the scale of measurement (community vs. plant). In addition, timing of grazing or defoliation emerged as a key factor that determined whether sexual reproduction was inhibited. Like the early season grazing that is typical of Yellowstone's northern winter range, studies often showed that early season defoliation stimulated production of community-level reproductive biomass. Our results rectify disagreements in the literature that ultimately derive from differences in either timing of defoliation or measurement scale. DOI:10.2458/azu_jrm_v56i5_anderson

Large-scale effects of a small herbivore on salt-marsh vegetation succession – A comparative study on three Wadden Sea islands

Grazing by livestock is used as a management tool to prevent the dominance of a single tall-growing specises during succession on European salt marshes. The effects of natural small herbivores are often neglected by managers. Long-term exclosure experiments on the island of Schiermonnikoog show that hares retard vegetation succession at the early stages of salt-marsh development. In the present study we test whether we can scale-up these exclosure studies to a whole salt-marsh system. We compared 30 years of undisturbed vegetation succession at the Wadden Sea islands of Schiermonnikoog, Rottumerplaat (both The Netherlands) and Mellum (Germany). Salt-marsh development started at all sites in the early 1970s. Hares have been present only on Schiermonnikoog. At each site an area was selected covering a gradient from high to low salt marsh. Surface elevation and clay thickness were measured and a vegetation map was made on the three islands. The areas showed similar clay thickness at low surface elevation, indicating similar sedimentation ratesand hence nitrogen inputs. Rottumerplaat and Mellum showed a higher dominance of the late successional speciesAtriplex portulacoides in the low marsh andElymus athericus in the high marsh compared to Schiermonnikoog. Typical mid-successional, important food plant species for hares and geese had a higher abundance at Schiermonnikoog. Patterns of vegetation development in the absence of hares followed the observed patterns in the smallscale exclosure experiments at Schiermonnikoog. Without hare grazing, vegetation succession proceeds more rapidly and leads to the dominance of tall-growing species in earlier stages of succession. The present study shows that next to large herbivores, small herbivores potentially have largescale effects on salt-marsh vegetation succession during the early successional stages.

Defoliation effects on reproductive biomass: importance of scale and timing

Community-level (per unit area) and individual tiller reproductive biomass inside and outside of long-term exclosures on the northern winter range of Yellowstone National Park, USA were compared. Grazed areas had twice the number of reproductive tillers m-2 (186 compared to 88 tillers m-2), and greater total reproductive biomass m-2 than ungrazed plots (13 compared to 7 g m-2). In contrast, seed number tiller-1 was greater for grasses in exclosures. Because of these offsetting responses, seed production (nom-2) was unaffected by herbivores. On an area basis, grazed grasses allocated proportionally more biomass to reproduction (reproductive biomass/aboveground biomass) than ungrazed grasses. We propose that altered plant demography and morphology following defoliation explain how grazers might increase the allocation of biomass to reproduction in Yellowstone grasslands. To understand these results in light of ecological and agronomic studies, we reviewed literature from 118 sources that reported the effects of defoliation on the production of reproductive biomass. The review suggested that the results of herbivory or defoliation on plant reproductive biomass depended on the scale of measurement (community vs. plant). In addition, timing of grazing or defoliation emerged as a key factor that determined whether sexual reproduction was inhibited. Like the early season grazing that is typical of Yellowstone's northern winter range, studies often showed that early season defoliation stimulated production of community-level reproductive biomass. Our results rectify disagreements in the literature that ultimately derive from differences in either timing of defoliation or measurement scale.

Grazing, Environmental Heterogeneity, and Alien Plant Invasions in Temperate Pampa Grasslands

Temperate humid grasslands are known to be particularly vulnerable to invasion by alien plant species when grazed by domestic livestock. The Flooding Pampa grasslands in eastern Argentina represent a well-documented case of a regional flora that has been extensively modified by anthropogenic disturbances and massive invasions over recent centuries. Here, we synthesise evidence from region-wide vegetation surveys and long-term exclosure experiments in the Flooding Pampa to examine the response of exotic and native plant richness to environmental heterogeneity, and to evaluate grazing effects on species composition and diversity at landscape and local community scales. Total plant richness showed a unimodal distribution along a composite stress/fertility gradient ranging several plant community types. On average, more exotic species occurred in intermediate fertility habitats that also contained the highest richness of resident native plants. Exotic plant richness was thus positively correlated with native species richness across a broad range of flood-prone grasslands. The notion that native plant diversity decreases invasibility was supported only for a limited range of species-rich communities in habitats where soil salinity stress and flooding were unimportant. We found that grazing promoted exotic plant invasions and generally enhanced community richness, whereas it reduced the compositional and functional heterogeneity of vegetation at the landscape scale. Hence, grazing effects on plant heterogeneity were scale-dependent. In addition, our results show that environmental fluctuations and physical disturbances such as large floods in the pampas may constrain, rather than encourage, exotic species in grazed grasslands.

EFFECTS OF MIGRATORY GRAZERS ON SPATIAL HETEROGENEITY OF SOIL NITROGEN PROPERTIES IN A GRASSLAND ECOSYSTEM

Large herbivores are known to respond to spatial heterogeneity in the distribution of forage resources, but how important herbivores are in creating those spatial patterns, how their influence may be scale dependent, and how herbivore-induced patterns affect ecosystem processes remain unclear. We examined the effects of native migratory ungulates on the spatial distribution of soil nitrogen and N-mineralization potential at scales ranging from 0.1–30 m in grasslands of Yellowstone National Park using geostatistical analyses of soils collected inside and outside long-term (36+-yr) exclosures. At small spatial scales (0.1–2 m), ungrazed grasslands showed a high degree of patchiness in the distribution of soil N and N-mineralization potential (94% and 77% of sample variation explained by small-scale patchiness, respectively). For both variables, patches occurred at a consistent mean size of ∼40 cm. In contrast, grazed grassland exhibited minimal spatial structure in the distribution of soil N and N-mineralization rates (<24% of variation for both variables spatially dependent) and no consistent patch size at a scale of 0.1–2 m. In grazed grassland, most variation was extremely fine grained, occurring across distances <10 cm. The high degree of fine-grained heterogeneity in grazed grassland was associated with greater plant diversity in small (20 × 20 cm) patches. Recycling of nitrogen through dung and urine is clearly important for maintaining long-term nitrogen balance of the system, but we estimated that only 2.5% of the grazed grassland sampled was recently affected by urine. Results conflict with predictions of increased heterogeneity in grazed communities based primarily on the deposition of dung and urine in discrete patches and suggest that the dominant mechanism(s) by which grazers alter N cycling in this ecosystem operates through the plant community. We hypothesize that grazers promote fine-scale heterogeneity by diversifying plant species effects on soils and/or increasing the spatial variability in plant litter inputs. At larger spatial scales (5–30 m), large herbivores altered the distribution of soil N across a topographic gradient. In a grazed community, soil N properties were associated with the topographic gradient and exhibited increasing variance among sampling points separated by increasing distances from 5 to 30 m. Ungrazed grassland exhibited no spatial structure in soil N distribution and no correlation with topography. Collectively, our data show that grazers influence the distribution of soil N properties at every spatial scale from individual plants to landscapes.

Changes induced by elk browsing in the aboveground biomass production and distribution of willow (Salix monticola Bebb): their relationships with plant water, carbon, and nitrogen dynamics.

Willows are dominant woody plants of many high-elevation riparian areas of the western USA, and constitute an important food resource for various ungulates, which tend to concentrate in riparian areas. The response of willow to browsing was analyzed in the elk winter range of Rocky Mountain National Park, by considering the effect of elk browsing on Salix monticola Bebb, one of the most common willow species in this area. Unbrowsed and browsed treatments were established during the 1997 growing season (May to October), using eight long-term exclosures built in the fall of 1994. Plants in the browsed treatment were in the areas open to browsing, but they were protected from browsing by small exclosures during the experimental period. Winter browsing by elk induced the following measured responses in plant morphology and development: (1) higher shoot biomass production but similar leaf biomass and leaf area per plant, (2) a lower number of and bigger shoots, (3) a lower number of and bigger leaves, and (4) flower inhibition. In addition, we infer that browsing induces (5) lower belowground allocation and (6) a more negative N balance but a higher soil N uptake. We conclude that elk browsing negatively affects willow even though willow compensate for aboveground biomass removal. Continuous browsing produces long-term changes in willow morphology which constrain plant growth and development. High browsing utilization, as occurred in this experiment, could therefore reduce the competitive ability and survivorship of willow, in particular under drier environmental conditions.