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.     

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.     

Evaluation of Central North American Prairie Management Based on Species Diversity, Life Form, and Individual Species Metrics

Abstract:  Reintroduction of fire and grazing, alone or in combination, has increasingly been recognized as central to the restoration of North American mixed-grass and tallgrass prairies. Although ecological studies of these systems are abundant, they have generally been observational, or if experimental, have focused on plant species diversity. Species diversity measures alone are not sufficient to inform management, which often has goals associated with life-form groups and individual species. We examined the effects of prescribed fire, light cattle grazing, and a combination of fire and grazing on three vegetation components: species diversity, groups of species categorized by life-form, and individual species. We evaluated how successful these three treatments were in achieving specific management goals for prairies in the Iowa Loess Hills (U.S.A.). The grazing treatment promoted the greatest overall species richness, whereas grazing and burning and grazing treatments resulted in the lowest cover by woody species. Burning alone best achieved the management goals of increasing the cover and diversity of native species and reducing exotic forb and (predominantly exotic) cool-season grass cover. Species-specific responses to treatments appeared idiosyncratic (i.e., within each treatment there existed a set of species attaining their highest frequency) and nearly half of uncommon species were present in only one treatment. Because all management goals were not achieved by any one treatment, we conclude that management in this region may need refining. We suggest that a mosaic of burning and grazing (alone and in combination) may provide the greatest landscape-level species richness; however, this strategy would also likely promote the persistence of exotic species. Our results support the need to consider multiple measures, including species-specific responses, when planning and evaluating management .     

High richness and dense seeding enhance grassland restoration establishment but have little effect on drought response

Restorations commonly utilize seed addition to formerly arable lands where the development of native plant communities is severely dispersal limited. However, variation in seed addition practices may profoundly affect restoration outcomes. Theory and observations predict that species-rich seed mixtures and seeding at high densities should enhance native plant community establishment, minimize exotic species cover, and may promote resistance and resilience to, and recovery from, environmental perturbations. We studied the post-seeding establishment of native plant communities in large grassland restoration plots, which were sown at two densities crossed with two levels of species richness on formerly arable land in Nebraska, USA, and their responses to drought. To evaluate drought resistance, recovery, and resilience of restored plant communities, we erected rainfall manipulation structures and tracked the response of seeded species cover and total plant biomass during experimental drought relative to controls and in the post-drought growing season. High seed richness and high-density seeding treatments resulted in greater richness and cover of native, seeded species per 0.5 m2 compared to low-richness and low-density treatments. Cover differences in response to seed mixture richness were driven by native forbs. Richness and cover of exotic species were lowest in high-richness and high-density treatments. We found little evidence of differential drought resistance, recovery, and resilience among seeding treatments. Increases in exotic species across years were restricted to drought subplots, and were not affected by seeding treatments. Grassland restoration was generally enhanced and exotic cover reduced both by the use of high-richness seed mixtures and high-density seeding. Given the lack of restoration treatment effects on the resistance, recovery, or resilience of seeded species exposed to drought, and the increases in exotic species following drought, other forms of active management may be needed to produce restored plant communities that are robust to climate change.     

Seed and establishment limitation contribute to long-term native forb declines in California grasslands

The effects of exotic species invasions on biodiversity vary with spatial scale, and documentation of local-scale changes in biodiversity following invasion is generally lacking. Coupling long-term observations of local community dynamics with experiments to determine the role played by exotic species in recruitment limitation of native species would inform both our understanding of exotic impacts on natives at local scales and regional-scale management efforts to promote native persistence. We used field experimentation to quantify propagule and establishment limitation in a suite of native annual forbs in a California reserve, and compared these findings to species abundance trends within the same sites over the past 48 years. Observations at 11 paired sites (inside and outside the reserve) indicated that exotic annual plants have continued to increase in abundance over the past 48 years. This trend suggests the system has not reached equilibrium > 250 years after exotic species began to spread, and 70 years after livestock grazing ceased within the reserve. Long-term monitoring observations also indicated that six native annual forb species went extinct from more local populations than were colonized. To determine the potential role of exotic species in these native plant declines, we added seed of these species into plots adjacent to monitoring sites where plant litter and live grass competition were removed. Experimental results suggest both propagule and establishment limitation have contributed to local declines observed for these native forbs. Recruitment was highest at sites that had current or historical occurrences of the seeded species, and in plots where litter was removed. Grazing history (i.e., location within or outside the reserve) interacted with exotic competition removal, such that removal of live grass competition increased recruitment in more recently grazed sites. Abundance of forbs was positively related to recruitment, while abundance of exotic forbs was negatively related. Thus, exotic competition is likely only one factor contributing to local declines of native species in invaded ecosystems, with a combination of propagule limitation, site quality, and land use history also playing important and interactive roles in native plant recruitment.     

Grassland Bird Responses to Land Management in the Largest Remaining Tallgrass Prairie

Abstract: Extensive habitat loss and changing agricultural practices have caused widespread declines in grassland birds throughout North America. The Flint Hills of Kansas and Oklahoma—the largest remaining tallgrass prairie—is important for grassland bird conservation despite supporting a major cattle industry. In 2004 and 2005, we assessed the community, population, and demographic responses of grassland birds to the predominant management practices (grazing, burning, and haying) of the region, including grasslands restored under the Conservation Reserve Program (CRP). We targeted 3 species at the core of this avian community: the Dickcissel (Spiza americana), Grasshopper Sparrow (Ammodramus savannarum), and Eastern Meadowlark (Sturnella magna). Bird diversity was higher in native prairie hayfields and grazed pastures than CRP fields, which were dominated by Dickcissels. Although Dickcissel density was highest in CRP, their nest success was highest and nest parasitism by Brown‐headed Cowbirds (Moluthrus ater) lowest in unburned hayfields (in 2004). Conversely, Grasshopper Sparrow density was highest in grazed pastures, but their nest success was lowest in these pastures and highest in burned hayfields, where cowbird parasitism was also lowest (in 2004). Management did not influence density and nest survival of Eastern Meadowlarks, which were uniformly low across the region. Nest success was extremely low (5–12%) for all 3 species in 2005, perhaps because of a record spring drought. Although the CRP has benefited grassland birds in agricultural landscapes, these areas may have lower habitat value in the context of native prairie. Hayfields may provide beneficial habitat for some grassland birds in the Flint Hills because they are mowed later in the breeding season than elsewhere in the Midwest. Widespread grazing and annual burning have homogenized habitat—and thus grassland‐bird responses—across the Flint Hills. Diversification of management practices could increase habitat heterogeneity and enhance the conservation potential of the Flint Hills for grassland birds.     

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.     

Cars, Cows, and Checkerspot Butterflies: Nitrogen Deposition and Management of Nutrient-Poor Grasslands for a Threatened Species

Abstract: Nutrient-poor, serpentinitic soils in the San Francisco Bay area sustain a native grassland that supports many rare species, including the Bay checkerspot butterfly ( Euphydryas editha bayensis ). Nitrogen (N) deposition from air pollution threatens biodiversity in these grasslands because N is the primary limiting nutrient for plant growth on serpentinitic soils. I investigated the role of N deposition through surveys of butterfly and plant populations across different grazing regimes, by literature review, and with estimates of N deposition in the region. Several populations of the butterfly in south San Jose crashed following the cessation of cattle grazing. Nearby populations under continued grazing did not suffer similar declines. The immediate cause of the population crashes was rapid invasion by introduced annual grasses that crowded out the larval host plants of the butterfly. Ungrazed serpentinitic grasslands on the San Francisco Peninsula have largely resisted grass invasions for nearly four decades. Several lines of evidence indicate that dry N deposition from smog is responsible for the observed grass invasion. Fertilization experiments have shown that soil N limits grass invasion in serpentinitic soils. Estimated N deposition rates in south San Jose grasslands are 10–15 kg  N/ha/year; Peninsula sites have lower deposition, 4–6 kg N/ha/year. Grazing cattle select grasses over forbs, and grazing leads to a net export of N as cattle are removed for slaughter. Although poorly managed cattle grazing can significantly disrupt native ecosystems, in this case moderate, well-managed grazing is essential for maintaining native biodiversity in the face of invasive species and exogenous inputs of N from nearby urban areas.     

Cover of Perennial Grasses in Southeastern Arizona in Relation to Livestock Grazing

Tolerance of particular grasslands to the activities of domestic livestock may depend on their historic association with native grazing animals. Southwestern grama ( Bouteloua ) grasslands are floristically allied to the North American Central Plains but lie outside the historic range of the plains' principal ungulate grazer, alics bishop . We compared perennial grassland cover and species composition on eight sites transacted by the boundary fence of a 3160-ha, 22-year-old livestock exclosure in a grama grassland in southeastern Arizona. Total grass canopy cover was greatest on the ungrazed portion of each of the eight sites. Two short stoloniferous species ( Hilaria belangeri and Bouteloua eriopoda ) were the only taxa substantially more abundant on grazed quadrats overall. Among these and eight taller budgerigars, there was a strong positive correlation between potential height and response to release from grazing, with the three tallest species showing the greatest increases on ungraded treatments ( emization curtailment, Boilermaker barbarians , and emizations intermixed ). emization gracious , the most abundant grass in the region, showed an intermediate response to livestock exclusion, Gram grasslands at the Arizona site have changed more and in different ways following livestock exclusion than those on the Central Plains of Colorado. Contributing factors may include: (1) greater annual precipitation at the Arizona site, (2) the much larger size of the Arizona livestock exclosure, and (3) the absence of extensive grazing by native ungulates in the Southwest since the Pleistocene. Livestock grazing appears to be an exotic ecological force in these southwestern grasslands, and one destructive of certain components of the native flora and fauna.     

Influence of a Large Herbivore Reintroduction on Plant Invasions and Community Composition in a California Grassland

Abstract:  Despite many successful reintroductions of large mammalian herbivores throughout the world, remarkably little attention has focused on how these actions affect native and exotic vegetation at reintroduction sites. One such herbivore is tule elk ( Cervus elaphus nannodes ), which was on the brink of extinction in the mid 1800s, but now has numerous stable populations due to intensive reintroduction efforts. Here, we summarize results from a 5-year exclosure experiment that explored the effects of tule elk on a coastal grassland in northern California. Elk significantly altered the species composition of this community; the response of annual species (dominated heavily by exotic taxa) was dramatically different from perennial species. Elk herbivory increased the abundance and aboveground biomass of native and exotic annuals, whereas it either had no effect on or caused significant decreases in perennials. Elk also decreased the cover of native shrubs, suggesting that these herbivores play an important role in maintaining open grasslands. In addition, elk significantly reduced the abundance and biomass of a highly invasive exotic grass , Holcus lanatus, which is a major problem in mesic perennial grasslands. Our results demonstrate that the successful reintroduction of a charismatic and long-extirpated mammal had extremely complex effects on the plant community, giving rise to both desirable and undesirable outcomes from a management perspective. We suspect that these kinds of opposing effects are not unique to tule elk and that land managers will frequently encounter them when dealing with reintroduced mammals.     

Direct Effects of Cattle on Grassland Birds in Canada

Effects of grazing on grassland birds are generally thought to be indirect, through alteration of vegetation structure; however, livestock can also affect nest survival directly through trampling and other disturbances (e.g., livestock‐induced abandonment). We extracted data on nest fates from 18 grazing studies conducted in Canada. We used these data to assess rates of nest destruction by cattle among 9 ecoregions and between seasonal and rotational grazing systems. Overall, few nests were destroyed by cattle (average 1.5% of 9132 nests). Nest destruction was positively correlated with grazing pressure (i.e., stocking rate or grazing intensity), but nest survival was higher in more heavily grazed areas for some species. Because rates of destruction of grassland bird nests by cattle are low in Canada, management efforts to reduce such destruction may not be of ecological or economic value in Canada. Efectos Directos del Ganado sobre las Aves de Pastizales en Canadá     

围封6年对温带典型草原坡地物种多样性及其与地上生物量的关系的影响

以内蒙古太仆寺旗地区典型克氏针茅-羊草草原为研究对象,通过调查围封6年后围栏内外典型草原群落特征,分析围封、放牧处理下典型草原植物群落结构及地上生物量的动态变化,揭示围封放牧对植被群落结构、物种丰富度和地上生物量的影响。结果发现,在围封6年后,各功能群物种丰富度与盖度围栏内外的空间变化趋势基本一致。群落物种丰富度和地上生物量均随坡位下降而显著增加,初步显示了物种多样性与生态系统功能的空间变化的一致性。围栏内物种丰富度显著低于围栏外,而地上生物量却显著大于围栏外。围栏内多年生禾草、一年生植物物种丰富度显著低于围栏外,而半灌木盖度、生物量显著高于围栏外。围栏内外在禁牧和放牧不同处理下,表现出不同的草原退化方式：在禁牧条件下,尽管生物量有所恢复,但灌木和半灌木的增加却代表了草原的退化;放牧、刈割等人类活动增加了耐牧物种和不适口植物生长的可能,表现为草原退化指示物种增加,同样是草原退化的表现。围栏外一年生植物盖度和生物量显著高于围栏内,而一年生植物的频数也远远大于围栏内（围栏内外频数比13：71）,且独行菜、猪毛菜等仅出现于围栏外。同时,地上生物量随着物种丰富度的增加而增加,但只有围栏外的关系达到统计显著程度,说明围封调制了生物多样性与生态系统功能的关系。本研究表明,对于干旱半干旱典型草原的恢复,不能简单地采用长期完全禁牧措施,应当根据区域环境、植被以及社会经济情况,制定季节性放牧或者间歇性禁牧的恢复措施,从而保证草场的可持续性恢复利用。     

Remote analysis of biological invasion and the impact of enemy release

Escape from natural enemies is a widely held generalization for the success of exotic plants. We conducted a large-scale experiment in Hawaii (USA) to quantify impacts of ungulate removal on plant growth and performance, and to test whether elimination of an exotic generalist herbivore facilitated exotic success. Assessment of impacted and control sites before and after ungulate exclusion using airborne imaging spectroscopy and LiDAR, time series satellite observations, and ground-based field studies over nine years indicated that removal of generalist herbivores facilitated exotic success, but the abundance of native species was unchanged. Vegetation cover <1 m in height increased in ungulate-free areas from 48.7% +/- 1.5% to 74.3% +/- 1.8% over 8.4 years, corresponding to an annualized growth rate of lambda = 1.05 +/- 0.01 yr(-1) (median +/- SD). Most of the change was attributable to exotic plant species, which increased from 24.4% +/- 1.4% to 49.1% +/- 2.0%, (lambda = 1.08 +/- 0.01 yr(-1)). Native plants experienced no significant change in cover (23.0% +/- 1.3% to 24.2% +/- 1.8%, lambda = 1.01 +/- 0.01 yr(-1)). Time series of satellite phenology were indistinguishable between the treatment and a 3.0-km2 control site for four years prior to ungulate removal, but they diverged immediately following exclusion of ungulates. Comparison of monthly EVI means before and after ungulate exclusion and between the managed and control areas indicates that EVI strongly increased in the managed area after ungulate exclusion. Field studies and airborne analyses show that the dominant invader was Senecio madagascariensis, an invasive annual forb that increased from < 0.01% to 14.7% fractional cover in ungulate-free areas (lambda = 1.89 +/- 0.34 yr(-1)), but which was nearly absent from the control site. A combination of canopy LAI, water, and fractional cover were expressed in satellite EVI time series and indicate that the invaded region maintained greenness during drought conditions. These findings demonstrate that enemy release from generalist herbivores can facilitate exotic success and suggest a plausible mechanism by which invasion occurred. They also show how novel remote-sensing technology can be integrated with conservation and management to help address exotic plant invasions.     

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.     

Bottom-up control of consumers leads to top-down indirect facilitation of invasive annual herbs in semiarid Chile

The abundance of exotic plants is thought to be limited by competition with resident species (including plants and generalist herbivores). In contrast, observations in semiarid Chile suggest that a native generalist rodent, the degu (Octodon degus), may be facilitating the expansion of exotic annual plants. We tested this hypothesis with a 20-year data set from a World Biosphere Reserve in mediterranean Chile. In this semiarid environment, rainfall varies annually and dramatically influences cover by both native and exotic annual plants; degu population density affects the composition and cover of exotic and native annual plants. In low-rainfall years, cover of both native and exotic herbs is extremely low. Higher levels of precipitation result in proportional increases in cover of all annual plants (exotic and native species), leading in turn to increases in degu population densities, at which point they impact native herbs in proportion to their greater cover, indirectly favoring the expansion of exotic plants. We propose that bottom-up control of consumers at our site results in top-down indirect facilitation of invasive annual herbs, and that this pattern may be general to other semiarid ecosystems.     

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.     

Postfire Management on Forested Public Lands of the Western United States

Abstract:  Forest ecosystems in the western United States evolved over many millennia in response to disturbances such as wildfires. Land use and management practices have altered these ecosystems, however, including fire regimes in some areas. Forest ecosystems are especially vulnerable to postfire management practices because such practices may influence forest dynamics and aquatic systems for decades to centuries. Thus, there is an increasing need to evaluate the effect of postfire treatments from the perspective of ecosystem recovery. We examined, via the published literature and our collective experience, the ecological effects of some common postfire treatments. Based on this examination, promising postfire restoration measures include retention of large trees, rehabilitation of firelines and roads, and, in some cases, planting of native species. The following practices are generally inconsistent with efforts to restore ecosystem functions after fire: seeding exotic species, livestock grazing, placement of physical structures in and near stream channels, ground-based postfire logging, removal of large trees, and road construction. Practices that adversely affect soil integrity, persistence or recovery of native species, riparian functions, or water quality generally impede ecological recovery after fire. Although research provides a basis for evaluating the efficacy of postfire treatments, there is a continuing need to increase our understanding of the effects of such treatments within the context of societal and ecological goals for forested public lands of the western United States.     

Experimental tree removal in tallgrass prairie: variable responses of flora and fauna along a woody cover gradient

Woody plant encroachment is a worldwide phenomenon in grassland and savanna systems whose consequence is often the development of an alternate woodland state. Theoretically, an alternate state may be associated with changes in system state variables (e.g., species composition) or abiotic parameter shifts (e.g., nutrient availability). When state-variable changes are cumulative, such as in woody plant encroachment, the probability of parameter shifts increases as system feedbacks intensify over time. Using a Before-After Control-Impact (BACI) design, we studied eight pairs of grassland sites undergoing various levels of eastern redcedar (Juniperus virginiana) encroachment to determine whether responses of flora and fauna to experimental redcedar removal differed according to the level of pretreatment redcedar cover. In the first year after removal, herbaceous plant species diversity and evenness, woody plant evenness, and invertebrate family richness increased linearly with pretreatment redcedar cover, whereas increases in small-mammal diversity and evenness were described by logarithmic trends. In contrast, increases in woody plant diversity and total biomass of terrestrial invertebrates were accentuated at levels of higher pretreatment cover. Tree removal also shifted small-mammal species composition toward a more grassland-associated assemblage. During the second year postremoval, increases in herbaceous plant diversity followed a polynomial trend, but increases in most other metrics did not vary along the pretreatment cover gradient. These changes were accompanied by extremely high growing-season precipitation, which may have homogenized floral and faunal responses to removal. Our results demonstrate that tree removal increases important community metrics among grassland flora and fauna within two years, with some responses to removal being strongly influenced by the stage of initial encroachment and modulated by climatic variability. Our results underscore the importance of decisive management for reversing the effects of woody plant encroachment in imperiled grassland ecosystems.     

Indirect Effects of Biological Control on Plant Diversity Vary across Sites in Montana Grasslands

Abstract:  Biological control with specialist, nonindigenous, herbivorous insects is an important option for controlling invasive exotic plants in wildlands and nature reserves. It is assumed that biological control agents will reduce the dominance of the target weed, thereby increasing the native diversity of the associated plant community. However, this hypothesis has rarely been tested. We introduced Aphthona nigriscutis into grassland sites infested with the invasive exotic species Euphorbia esula L. on a nature reserve in Montana (U.S.A.). Two sites with better soil had been treated previously with herbicide, whereas two other sites had not. We measured the density and biomass of Euphorbia vegetative and flowering stems and number of native and exotic shrubs, grass-like plants, and forbs in 48 microplots in Aphthona release and control macroplots at each site. After 5 years, Aphthona release was associated with a 33–39% decline in Euphorbia aboveground biomass compared with controls at all sites. Other effects of the biocontrol depended on the site. Biocontrol slowed the recovery of species diversity at the sites previously treated with herbicide but slowed the loss of diversity at sites without a history of herbicide. Biocontrol introduction was not associated with a disproportionate increase in nontarget exotic species. Release of Aphthona caused a decline in the biomass of flowering stems relative to controls at good-soil, previous-herbicide sites but was associated with a relative increase in flower stem mass at poor-soil, no-herbicide sites. Our results suggest that biocontrol reductions in weed dominance will not always be associated with increased species diversity. More emphasis should be placed on conserving desirable communities and less on simple weed control. Monitoring of community-level effects should accompany biocontrol introductions on nature reserves.     

Twenty years of salt marsh succession on a Dutch coastal barrier island

After a formerly grazed salt marsh was released from cattle grazing, changes in plant species composition were monitored for 20 yr, using vegetation maps and permanent plots. Three areas, differing in age and nutrient status were compared. The number of plant species and plant communities decreased.Elymus athericus (Elytrigia pungens) became dominant in most plant communities after 5–20 yr on the oldest and most productive salt marsh. In younger areas it took more time forE. athericus to become dominant. At least 7 cm of clay seemed to be a prerequisite for this plant species to increase in dominance. The results from monitoring over decades are discussed in view of the knowledge on succession over centuries as derived from a chronosequence.