Cattle Grazing Impacts on Annual Forbs and Vegetation Composition of Mesic Grasslands in California

Abstract:   Livestock grazing represents a major human alteration of natural disturbance regimes in grasslands throughout the world, and its impacts on plant communities have been highly debated. We investigated the impact of cattle grazing on the California coastal prairie plant community with a focus on native annual forbs, a number of which are of conservation concern. In spring 2000 and 2001, we surveyed the vegetation community composition, vegetation structure, and soil chemical parameters at 25 paired grazed and ungrazed sites over a 670-km range of the ecosystem. Native annual forb species richness and cover were higher in grazed sites, and this effect was concomitant with decreased vegetation height and litter depth. Soil properties explained less of the variation. Exotic annual grass and forb cover were higher in grazed sites. Native grass cover and species richness did not differ in grazed and ungrazed sites, but cover and species richness of native perennial forbs were higher in ungrazed sites. Our results suggest that cattle grazing may be a valuable management tool with which to conserve native annual forbs in the ecosystem we studied but that grazing differentially affects the various life-history guilds. Therefore, land managers must focus on creating a matrix of disturbance regimes to maintain the suite of species native to these mesic grasslands. The results of this and other studies highlight the importance of considering the adaptation of vegetation communities to disturbance in making recommendations for grazing management.     

Short-term and long-term facilitation of goose grazing by livestock in the Dutch Wadden Sea area

We studied the impact of livestock grazing on the distribution ofBranta bernicla bernicla (Dark-bellied Brent goose) in the Dutch Wadden Sea during spring. It was hypothesized that livestock facilitate short-term (within-season) grazing for geese as well as long-term (over years). Therefore we measured grazing pressure by geese in salt marsh and polder areas that were either grazed (spring-grazed) or ungrazed during spring (summer-grazed). Additionally, we carried out a preference experiment with captive geese to test the preference between spring-grazed and summer-grazed polder swards. We furthermore compared patterns of use by geese between long-term ungrazed and grazed salt marshes. In May, there is a difference in grazing pressure by geese between polder pastures that are grazed or ungrazed during spring. In this month, the ungrazed polder pastures are abandoned and the geese shift to either the grazed polder pastures or to the salt marsh. Vegetation in the polder that had been spring-grazed had a lower canopy height and a higher tiller density than summer-grazed vegetation. The captive geese in the preference experiment showed a clear preference for vegetation that had been spring-grazed by sheep over ungrazed vegetation. Goose grazing pressure was negatively correlated to canopy height, both on the polder and on the salt marsh. Within the plant communities dominated byFestuca rubra andPuccinellia maritima, marshes that were intensively grazed by livestock generally had higher grazing pressure by geese than long-term ungrazed or lightly grazed salt marshes.     

The impact of grazing on spider communities in a mesophytic calcareous dune grassland

During 1994–1995 and 1997–1998 spiders were sampled with pitfall traps in a botanically rich, mesophytic, calcareous dune grassland in Belgium. As a consequence of intensive cattle grazing, vegetation variation in a large part of the area had diminished. The study area was also patchily grazed by rabbits. Community analysis with TWINSPAN revealed five distinct spider communities. Ecological differentiation was best explained by combination of the habitat variables: distance from grazed or non-grazed vegetation,Rosa pimpinellifolia cover and grass cover in both summer and winter. Species diversity was highest in the border zone between the cattle-grazed and non cattle-grazed sites. Correlation of the most abundant spider species with the vegetation determinants explains the ecological differentiation between the spider communities. Species were classified into seven major groups that reflect the species’ habitat preferences. The group showing clear association with non cattle-grazed, tall vegetation consists of common species. Characteristic species for the intensively cattle-grazed sites are common aeronauts and rare species such asWalckenaeria stylifrons, Mastigusa arietina, Ceratinopsis romana andPardosa monticola. The latter are shown to be dependent on ungrazed vegetation for juvenile development and overwintering. Intensive grazing results in homogeneous short vegetation, which can only be colonized by ‘open ground’ species with a well-developed dispersal capacity, or by species which are not dependent on litter-rich situations for juvenile development. An extensive cattle grazing regime results in a patchy mosaic grassland where, in addition to the above mentioned groups of species, other species survive by migrating between the buffered litter rich ungrazed vegetation and the short vegetation. Additionally, some typical and rare species prefer the transition zone between the grazed and the ungrazed vegetation because they are associated with specific habitat structures or inhabiting ant-species.     

Effects of Cattle Grazing on Diversity in Ephemeral Wetlands

Abstract:  Cattle are usually thought of as a threat to biodiversity. In regions threatened by exotic species invasion and lacking native wild grazers, however, cattle may produce the type of disturbance that helps maintain diverse communities. Across 72 vernal pools, I examined the effect of different grazing treatments (ungrazed, continuously grazed, wet-season grazed and dry-season grazed) on vernal-pool plant and aquatic faunal diversity in the Central Valley of California. After 3 years of treatment, ungrazed pools had 88% higher cover of exotic annual grasses and 47% lower relative cover of native species than pools grazed at historical levels (continuously grazed). Species richness of native plants declined by 25% and aquatic invertebrate richness was 28% lower in the ungrazed compared with the continuously grazed treatments. Release from grazing reduced pool inundation period by 50 to 80%, making it difficult for some vernal-pool endemic species to complete their life cycle. My results show that one should not assume livestock and ranching operations are necessarily damaging to native communities. In my central California study site, grazing helped maintain native plant and aquatic diversity in vernal pools.     

The impact of grazing on spider communities in a mesophytic calcareous dune grassland

During 1994–1995 and 1997–1998 spiders were sampled with pitfall traps in a botanically rich, mesophytic, calcareous dune grassland in Belgium. As a consequence of intensive cattle grazing, vegetation variation in a large part of the area had diminished. The study area was also patchily grazed by rabbits. Community analysis with TWINSPAN revealed five distinct spider communities. Ecological differentiation was best explained by combination of the habitat variables: distance from grazed or non-grazed vegetation,Rosa pimpinellifolia cover and grass cover in both summer and winter. Species diversity was highest in the border zone between the cattle-grazed and non cattle-grazed sites. Correlation of the most abundant spider species with the vegetation determinants explains the ecological differentiation between the spider communities. Species were classified into seven major groups that reflect the species’ habitat preferences. The group showing clear association with non cattle-grazed, tall vegetation consists of common species. Characteristic species for the intensively cattle-grazed sites are common aeronauts and rare species such asWalckenaeria stylifrons, Mastigusa arietina, Ceratinopsis romana andPardosa monticola. The latter are shown to be dependent on ungrazed vegetation for juvenile development and overwintering. Intensive grazing results in homogeneous short vegetation, which can only be colonized by ‘open ground’ species with a well-developed dispersal capacity, or by species which are not dependent on litter-rich situations for juvenile development. An extensive cattle grazing regime results in a patchy mosaic grassland where, in addition to the above mentioned groups of species, other species survive by migrating between the buffered litter rich ungrazed vegetation and the short vegetation. Additionally, some typical and rare species prefer the transition zone between the grazed and the ungrazed vegetation because they are associated with specific habitat structures or inhabiting ant-species. Nomenclature: Roberts (1987, 1995) forAraneae; van der Meijden et al. (1990) for vascular plants; Corly et al. (1981) for bryophytes; Schaminée et al. (1996) for vegetation associations.     

Effect of Vertebrate Grazing on Plant and Insect Community Structure

Abstract: We compared species diversity of plants and insects among grazed and ungrazed areas of Ponderosa pine–grassland communities in Arizona. Plant species richness was higher in two of three grassland communities that were grazed by native elk and deer and domestic cattle than in ungrazed areas inside a series of three large (approximately 40-ha) grazing exclosures. Similarly, plant species richness was higher in grazed areas relative to ungrazed areas at one of two series of smaller (approximately 25-m²) and short-term exclosure sites. Evenness of plant distribution, however, was greater inside ungrazed long-term exclosures but was reduced inside ungrazed short-term exclosures relative to grazed areas. Relative abundances of forbs, grasses, trees, and shrubs, and native and introduced plants did not differ between the long- and short-term grazing exclosures and their grazed counterparts. Relative abundances of some plant species changed when grazers were excluded, however. In contrast, insect species richness was not different between grazed and ungrazed habitats, although insect abundance increased 4- to 10-fold in ungrazed vegetation. Our results suggest that vertebrate grazing may increase plant richness, even in nutrient-poor, semi-arid grasslands, but may decrease insect abundances.     

Grazing as a measure against grass encroachment in Dutch dry dune grassland: effects on vegetation and soil

A grazing experiment was started in 1984 and 1989 respectively, in two parts of a dune grassland in the nature reserve ‘Zwanenwater’, North Holland; a third part with similar geology and topography was used as a control area and not grazed. An evaluation of the effects of grazing on vegetation patterns, species composition, vegetation structure and humus form was made with the help of vegetation maps from 1986 and 1992 as well as field surveys. Dense tall-grass communities dominated byAmmophila arenaria increased over the period 1986–1992 in the grazed areas, and especially in the non-grazed area (increase in area to 20%, 22% and 50%, respectively). Open communities decreased in the grazed areas, but are still prevalent, while in the ungrazed area they virtually disappeared, with the result that the present percentage areas are 53%, 38% and 17%. Field survey data were classified by TWINSPAN producing four vegetation types. These occur more or less equally in grazed and ungrazed areas, albeit with different percentage areas: (1) open vegetation dominated byCorynephorus canescens; (2) open vegetation characteized, byKoeleria macrantha; (3) heathland dominated byEmpetrum nigrum; and (4) tall-grass communities dominated byAmmophila arenaria. Within a vegetation type, species composition was only marginally affected by grazing regime. Within the open communities the number of species, vegetation height, vegetation cover and soil organic horizons were not affected by grazing. In the tall-grass communities the number of species was significantly larger and the height of the vegetation significantly lower in the area grazed since 1984. In the heathland community the number of species and cover of the moss layer were significantly higher in the 1984 area and ectorganic and endorganic horizons significantly thicker in the ungrazed area. It is suggested that these effects are the result of an increased availability of light, but possibly also of a decreased stock of organic matter and nutrients, due to a decreased input of litter and accelerated rates of decomposition.     

Vegetation development influenced by grazing in the coastal dunes near The Hague, The Netherlands

In 1990, grazing was introduced in a section of Meijendel, a coastal sand dune system near The Hague, The Netherlands. After five years an evaluation was made of the effects of grazing on vegetation development. Three transects were established, two in grazed areas and one in an ungrazed area. Field survey data were classified by means of TWIN-SPAN, ordinated with Detrended Correspondence Analysis and the resulting vegetation types interpreted according to Westhoff & den Held (1969). All associations were found in both the grazed and the ungrazed areas, but at the subassociation and variant level some communities appeared to be restricted to the grazed area. These variants were five grassland variants characterized by disturbance indicators such asSenecio sylvaticus andCynoglossum officinale. The total number of plant species in the 19 permanent plots, which had been observed to have been decreasing since 1960, showed a considerable increase after the introduction of horses and cows in 1990. A marked decrease in the cover ofCalamagrostis epigejos andCarex arenaria since 1990 was evident, while in some plots species such asRibes rubrum andViburnum opulus increased considerably. A series of false-colour aerial photographs were used to compare vegetation structure in the three transects between 1990 and 1995. In the grazed area the tall grass vegetation had almost totally disappeared, whereas the areas of open sand. sand with moss and lichens, and low grass vegetation had increased and the pattern had become more fine-grained. In the ungrazed area the area covered by low grass vegetation had increased at the expense of the area of sand with moss and lichens and the pattern had become more coarse-grained.     

Interactive effects of rabbit grazing and environmental factors on plant species-richness on dunes of Norderney

On the German island Norderney rabbit grazing effects on vegetation, soil and microclimate were investigated with exclosures in different dune vegetation. Differences in species composition between grazed and ungrazed plots increased with grazing activity. A decline in growth height resulted in higher light availability and temperature on the grazed sites. One of the most visible effects was the decline in flowering plants. Most grazing effects varied between habitat types. Grey dunes were distinctly affected by rabbit grazing. In relation to the habitat type, life forms were differently affected: in closed vegetation, rabbit activities further open patches, resulting in more therophytes. In contrast, in open vegetation rabbit activities were too strong and following sand dynamics too high for many therophytes. In general, grazing diversified habitats, slightly supported the total species-richness, and enhanced the heterogeneity of species composition. Highest species-richness in total and of herbs was reached at intermediate levels of grazing pressure. The study is in line that highest species-diversity will be found at intermediate levels of disturbance and environmental stress and that grazing effects vary with the habitat type.     

Grazing as a measure against grass encroachment in Dutch dry dune grassland: effects on vegetation and soil

A grazing experiment was started in 1984 and 1989 respectively, in two parts of a dune grassland in the nature reserve ‘Zwanenwater’, North Holland; a third part with similar geology and topography was used as a control area and not grazed. An evaluation of the effects of grazing on vegetation patterns, species composition, vegetation structure and humus form was made with the help of vegetation maps from 1986 and 1992 as well as field surveys. Dense tall-grass communities dominated byAmmophila arenaria increased over the period 1986–1992 in the grazed areas, and especially in the non-grazed area (increase in area to 20 %, 22 % and 50 %, respectively). Open communities decreased in the grazed areas, but are still prevalent, while in the ungrazed area they virtually disappeared, with the result that the present percentage areas are 53 %, 38 % and 17 %. Field survey data were classified by TWINSPAN producing four vegetation types. These occur more or less equally in grazed and ungrazed areas, albeit with different percentage areas: (1) open vegetation dominated byCorynephorus canescens; (2) open vegetation characterized byKoeleria macrantha; (3) heathland dominated byEmpetrum nigrum; and (4) tall-grass communities dominated byAmmophila arenaria. Within a vegetation type, species composition was only marginally affected by grazing regime. Within the open communities the number of species, vegetation height, vegetation cover and soil organic horizons were not affected by grazing. In the tall-grass communities the number of species was significantly larger and the height of the vegetation significantly lower in the area grazed since 1984. In the heathland community the number of species and cover of the moss layer were significantly higher in the 1984 area and ectorganic and endorganic horizons significantly thicker in the ungrazed area. It is suggested that these effects are the result of an increased availability of light, but possibly also of a decreased stock of organic matter and nutrients, due to a decreased input of litter and accelerated rates of decomposition. Nomenclature: Van der Meijden et al. (1992) for phanerogams; Corley et al. (1981) for mosses; Grolle (1983) for hepatics.     

A trophic cascade triggers collapse of a salt-marsh ecosystem with intensive recreational fishing

Overexploitation of predators has been linked to the collapse of a growing number of shallow-water marine ecosystems. However, salt-marsh ecosystems are often viewed and managed as systems controlled by physical processes, despite recent evidence for herbivore-driven die-off of marsh vegetation. Here we use field observations, experiments, and historical records at 14 sites to examine whether the recently reported die-off of northwestern Atlantic salt marshes is associated with the cascading effects of predator dynamics and intensive recreational fishing activity. We found that the localized depletion of top predators at sites accessible to recreational anglers has triggered the proliferation of herbivorous crabs, which in turn results in runaway consumption of marsh vegetation. This suggests that overfishing may be a general mechanism underlying the consumer-driven die-off of salt marshes spreading throughout the western Atlantic. Our findings support the emerging realization that consumers play a dominant role in regulating marine plant communities and can lead to ecosystem collapse when their impacts are amplified by human activities, including recreational fishing.     

Effects of Historic Livestock Grazing on Vegetation at Chaco Culture National Historic Park, New Mexico

Abstract:   Livestock grazing is the most ubiquitous land use in western North America, yet it rarely has been studied in a controlled manner because of the lack of large areas free of grazing. We compared the ecological effects of three grazing treatments—long-term protection, short-term protection, and currently grazed—at Chaco Culture National Historic Park in northern New Mexico. Chaco has a long history of human habitation and is now one of the largest grazing exclosures in the American West. We studied the effects of livestock grazing on the cover of plants, soil crusts, and plant species richness at six sites with different potential natural vegetation. Species richness was higher under long-term protection than under current grazing at all six sites. Trends in shrub and grass response varied significantly across the six sites. Shrub cover increased with long-term protection at four upland sites, and grass cover increased with protection at four sites. The response of Chaco vegetation to release from grazing varied significantly according to each site's ecological potential, determined in part by edaphic and topographic characteristics. These nuances in vegetation response represent natural ecological variation and contrast with the notions of widespread shrub "invasion" often inferred in the past.     

Utilisation of a coastal grassland by geese after managed re-alignment

In this study we evaluate the effect of coastal re-alignment on the utilisation of coastal grasslands by staging geese. We assessed vegetation change and utilisation by geese using repeated mapping and regular dropping counts in both the restored marsh and adjacent reference sites. All measurements were started well before the actual re-alignment. In addition, we studied the effects of livestock grazing on vegetation and geese, using exclosures. The vegetation transformed from fresh grassland into salt-marsh vegetation. A relatively large proportion of the de-embanked area became covered with secondary pioneer vegetation, and the overall cover of potential food plants for geese declined. Goose utilisation had initially dropped to low levels, both in autumn and in spring, but it recovered to a level comparable to the reference marsh after ten years. Exclosure experiments revealed that livestock grazing prevented the establishment of closed swards of grass in the poorly drained lower area of the restored marsh, and thereby negatively affected goose utilisation of these areas during spring staging. Goose grazing in the restored marsh during spring showed a positive numerical response to grass cover found during the preceding growing season. (1) The value of restored salt marsh as foraging habitat for geese initially decreased after managed re-alignment but recovered after ten years. (2) Our findings support the idea that the value of foraging habitats depends largely on the cover of forage plants and that this can be manipulated by adjusting both grazing and drainage.     

Secondary succession on a high salt marsh at different grazing intensities

Succession occurs on a large part of German salt marshes following abandonment or reduction of grazing. Its speed and effect on the biodiversity of salt marshes has been discussed in the literature. Permanent plot studies show site-dependent differences in successional outcome. If grazing is to be continued, there is uncertainty about the stocking rates that are optimal for the conservation of plant diversity. We present an eight year permanent plot study with different grazing treatments on a high salt marsh at the marsh island of Hallig Langeness, Germany, which is enclosed by a summer dike. The study site is owned by WWF. Successional outcomes on the permanent plots depend on grazing intensity and depth of the groundwater table which is correlated with soil salinity. Intermediate stocking rates support a mixture of halophytes and glycophytes, the relative proportion of which depends on the depth of the groundwater table. Cattle grazing at stocking rates of 0.6 livestock units per ha were found to be optimal for plant diversity conservation. At sites with deeper groundwater tables and no grazing, dominantElymus spp. stands develop and diversity is strongly reduced. If the groundwater table is high, succession following grazing abandonment is retarded and small halophytes prevail.     

Rodent Communities in an Exurbanizing Southwestern Landscape (U.S.A.)

Abstract:  Ranches are being converted to exurban housing developments in the southwestern United States, with potentially significant but little-studied impacts on biological diversity. We captured rodents on 48 traplines in grasslands, mesquite savannas, and oak savannas in southeastern Arizona that were grazed by livestock, embedded in exurban housing developments, grazed and embedded in development, or neither grazed nor embedded in development. Independent of habitat or development, rodent species richness, mean rank abundance, and capture rates of all rodents combined were negatively related to presence of livestock grazing or to its effects on vegetative ground cover. Exurban development had no obvious effects on rodent variety or abundance. Results suggest southwestern exurban developments can sustain a rich assemblage of grassland and savanna rodents if housing densities are low and houses are embedded in a matrix of natural vegetation with little grazing.     

Cattle Grazing Mediates Climate Change Impacts on Ephemeral Wetlands

Abstract:  Climate change impacts depend in large part on land-management decisions; interactions between global changes and local resource management, however, rarely have been quantified. We used a combination of experimental manipulations and simulation modeling to investigate the effects of interactions between cattle grazing and regional climate change on vernal pool communities. Data from a grazing exclosure study indicated that 3 years after the removal of grazing, ungrazed vernal pools dried an average of 50 days per year earlier than grazed control pools. Modeling showed that regional climate change could also alter vernal pool hydrology. Increased temperatures and winter precipitation were predicted to increase periods of inundation. We evaluated the ecological implications of interactions between grazing and climate change for branchiopods and the California tiger salamander (  Ambystoma californiense ) at four sites spanning a latitudinal climate gradient. Grazing played an important role in maintaining the suitability of vernal pool hydrological conditions for fairy shrimp and salamander reproduction. The ecological importance of the interaction varied nonlinearly across the region. Our results show that grazing can confound hydrologic changes driven by climate change and play a critical role in maintaining the hydrologic suitability of vernal pools for endangered aquatic invertebrates and amphibians. These observations suggest an important limitation of impact assessments of climate change based on experiments in unmanaged ecosystems. The biophysical impacts of land management may be critical for understanding the vulnerability of ecological systems to climate change.     

Effects of Grazing on the Demography and Growth of the Texas Tortoise

Abstract: Considerable effort has been exerted in attempts to understand the complex ecological effects of grazing. North American tortoises, by virtue of their distribution, provide a good model taxon through which to study how grazing effects vary with grazing regime, habitat, and climate. We studied the Texas tortoise (  Gopherus berlandieri ) , which is restricted primarily to privately owned rangelands of southern Texas and northeastern Mexico. Management of this species is hampered by a lack of information on the effects of common land-use practices. We evaluated the effects of moderate grazing by cattle (short-duration, winter-spring rotational grazing regime; 6–28 animal-unit days/ha/year) on this tortoise by comparing two grazed and two ungrazed sites in the Western Rio Grande Plains, Texas ( U.S.A.), from April 1994 to October 1997. We made 132 captures of 106 individuals in the ungrazed pastures and 324 captures of 237 individuals in the grazed pastures. We also radiotracked 22 tortoises in the ungrazed pastures and 25 tortoises in the grazed pastures. Comparisons of relative abundance, body-size distribution, age distribution, body mass, sex ratio, adult survival, proportion of juveniles, and growth rates revealed no differences (  p > 0.05 for all parameters) between tortoises on grazed and ungrazed areas. Based on these results, we suggest that moderate grazing by cattle is not incompatible with maintenance of Texas tortoise populations. Our data were consistent with a general model of tortoise biogeography and tolerance of disturbance which suggests that Texas tortoises are tolerant to intermediate levels of disturbance. Generalities about the effect of cattle grazing on the four North American tortoises should be avoided unless they can be placed in the context of grazing regime, precipitation, habitat quality, and tortoise requirements.     

A robust-design formulation of the incidence function model of metapopulation dynamics applied to two species of rails

The incidence function model (IFM) uses area and connectivity to predict metapopulation dynamics. However, false absences and missing data can lead to underestimates of the number of sites contributing to connectivity, resulting in overestimates of dispersal ability and turnovers (extinctions plus colonizations). We extend estimation methods for the IFM by using a hierarchical Bayesian model to account both for false absences due to imperfect detection and for missing data due to sites not surveyed in some years. We compare parameter estimates, measures of metapopulation dynamics, and forecasts using stochastic patch occupancy models (SPOMs) among three IFM models: (1) a Bayesian formulation assuming no false absences and omitting site-year combinations with missing data; (2) a hierarchical Bayesian formulation assuming no false absences but incorporating missing data; and (3) a hierarchical Bayesian formulation allowing for imperfect detection and incorporating missing data. We fit the models to multiyear data sets of occupancy for two bird species that differ in body size and presumed dispersal ability but inhabit the same network of sites: the small Black Rail (Laterallus jamaicensis) and the medium-sized Virginia Rail (Rallus limicola). Incorporating missing data affected colonization parameters and led to lower estimates of dispersal ability for the Black Rail. Detection rates were high for the Black Rail in most years but moderate for the Virginia Rail. Incorporating imperfect detection resulted in higher occupancy and lower turnover rates for both species, with largest effects for the Virginia Rail. Forecasts using SPOMs were sensitive to both missing data and false absences; persistence in models assuming no false absences was more optimistic than from robust models. Our results suggest that incorporating false absences and missing data into the IFM can improve (1) estimates of dispersal ability and the effect of connectivity on colonization, (2) the scaling of extinction risk with patch area, and (3) forecasts of occupancy and turnover rates.     

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.     

Differing Effects of Cattle Grazing on Native and Alien Plants

Abstract:   Habitat managers use cattle grazing to reduce alien plant cover and promote native species in California grasslands and elsewhere in the western United States. We tested the effectiveness of grazing as a restoration method by examining the effects of herbivory on native and alien plants. At Carrizo Plain National Monument, California, we surveyed native and alien species cover in adjacent grazed and ungrazed areas. We also established experimental plots in which plants were clipped or mulch (dead biomass) was removed. In addition, we clipped plants grown in pots and plants in the field that grew with and without competitors. Native species were negatively affected by clipping in 1999, 2000, and 2001, whereas alien species were unaffected. In the experimental field plots, the European annual forb Erodium cicutarium compensated in growth and reproduction following simulated herbivory. In contrast, growth and reproduction of the native perennial bunchgrass Poa secunda were reduced 1 year after clipping. In pots, E. cicutarium overcompensated and grasses undercompensated. In the field, European grasses were unaffected by the removal of competitors. It is unclear by what mechanism E. cicutarium was able to compensate, but the ability may be related to its basal rosette growth form and indeterminately growing inflorescences. The native California grassland community assembled in the absence of grazing herds, whereas invasive European species have been exposed to grazing for centuries. It may be that these invaders have adaptations that better enable them to recover from grazing. In the grassland we studied, the strategy of livestock grazing for restoration is counterproductive. It harms native species and promotes alien plant growth.