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.     

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.     

Shifts in positive and negative plant interactions along a grazing intensity gradient

Isolating the single effects and net balance of negative and positive species effects in complex interaction networks is a necessary step for understanding community dynamics. Facilitation and competition have both been found to operate in harsh environments, but their relative strength may be predicted to change along gradients of herbivory. Moreover, facilitation effects through habitat amelioration and protection from herbivory may act together determining the outcome of neighborhood plant-plant interactions. We tested the hypothesis that grazing pressure alters the balance of positive and negative interactions between palatable and unpalatable species by increasing the strength of positive indirect effects mediated by associational resistance to herbivory. We conducted a two-year factorial experiment in which distance (i.e., spatial association) from the nearest unpalatable neighbor (Stipa speciosa) and root competition were manipulated for two palatable grasses (Poa ligularis and Bromus pictus), at three levels of sheep grazing (none, moderate, and high) in a Patagonian steppe community. We found that grazing shifted the effect of Stipa on both palatable grasses, from negative (competition) in the absence of grazing to positive (facilitation) under increasing herbivore pressure. In ungrazed sites, belowground competition was the dominant interaction, as shown by a significant reduction in performance of palatable grasses transplanted near to Stipa tussocks. In grazed sites, biomass of palatable plants was greater near than far from Stipa regardless of competition treatment. Proximity to Stipa reduced the amount of herbivory suffered by palatable grasses, an indirect effect that was stronger under moderate than under intense grazing. Our results demonstrate that facilitation, resulting mainly from protection against herbivory, is the overriding effect produced by unpalatable neighbors on palatable grasses in this rangeland community. This finding challenges the common view that abiotic stress amelioration should be the predominant type of facilitation in arid environments and highlights the role of herbivory in modulating complex neighborhood plant interactions in grazing systems.     

The adaptive value of remnant native plants in invaded communities: an example from the Great Basin

Changes in the species composition of biotic communities may alter patterns of natural selection occurring within them. Native perennial grass species in the Intermountain West are experiencing a shift in the composition of interspecific competitors from primarily perennial species to an exotic, annual grass. Thus traits that confer an advantage to perennial grasses in the presence of novel annual competitors may evolve in invaded communities. Here I show that such traits are apparent in populations of a native perennial grass, big squirreltail (Elymus multisetus M.E. Jones), exposed to cheatgrass (Bromus tectorum L.) competitors. Dormant big squirreltail plants were collected from cheatgrass-invaded and uninvaded sites near Bordertown, California, USA, a mid-elevation (1600 m) sagebrush community, and transplanted into pots in a greenhouse. Individual plants were split into equal halves. One half was grown with competition from cheatgrass, and the other half was grown without competition. Plants collected from invaded sites responded more quickly to watering, growing more leaves in the first 10 days after transplanting. In addition, big squirreltail plants collected from invaded areas experienced a smaller decrease in plant size when grown with competition than did plants collected from uninvaded areas. Accordingly, while there were fewer big squirreltail individuals in the invaded sites, they were more competitive with cheatgrass than were the more abundant conspecifics in nearby uninvaded areas. It is possible that annual grasses were the selective force that caused these population differences, which may contribute to the long-term persistence of the native populations. While it is tempting to restore degraded areas to higher densities of natives (usually done by bringing in outside seed material), such actions may impede long-term adaptation to new conditions by arresting or reversing the direction of ongoing natural selection in the resident population. If hot spots of rapid evolutionary change can be identified within invaded systems, these areas should be managed to promote desirable change and could serve as possible sources of restoration material or reveal traits that should be prioritized during the development of restoration seed material.     

Impact of Grazing Intensity during Drought in an Arizona Grassland

Abstract:  The ecological benefits of changing cattle grazing practices in the western United States remain controversial, due in part to a lack of experimentation. In 1997 we initiated an experimental study of two rangeland alternatives, cattle removal and high-impact grazing, and compared grassland community responses with those with more conventional, moderate grazing practices. The study was conducted in a high-elevation, semiarid grassland near Flagstaff, Arizona (U.S.A.). We conducted annual plant surveys of modified Whittaker plots for 8 years and examined plant composition shifts among treatments and years. High-impact grazing had strong directional effects that led to a decline in perennial forb cover and an increase in annual plants, particularly the exotic cheatgrass ( Bromus tectorum L.). A twofold increase in plant cover by exotic species followed a severe drought in the sixth year of the study, and this increase was greatest in the high-impact grazing plots, where native cover declined by one-half. Cattle removal resulted in little increase in native plant cover and reduced plant species richness relative to the moderate grazing control. Our results suggest that some intermediate level of cattle grazing may maintain greater levels of native plant diversity than the alternatives of cattle removal or high-density, short-duration grazing practices. Furthermore, episodic drought interacts with cattle grazing, leading to infrequent, but biologically important shifts in plant communities. Our results demonstrate the importance of climatic variation in determining ecological effects of grazing practices, and we recommend improving conservation efforts in arid rangelands by developing management plans that anticipate this variation.     

Fungal endophytes directly increase the competitive effects of an invasive forb

Competitive outcomes among plants can vary in different abiotic and biotic conditions. Here we tested the effects of two phylotypes of Alternaria endophytes on the growth, competitive effects, and competitive responses of the exotic invasive forb Centaurea stoebe. Centaurea stoebe was a better competitor against North American grass species than grasses from its European home range in the absence of endophytes. However, one endophyte both increased the biomass of C. stoebe and reduced the competitive effect of North American grasses on C. stoebe. The competitive effects of C. stoebe on grass species native to North America were enhanced by both fungal endophytes, but not for native European grasses. We do not know the mechanism by which endophytes increased C. stoebe's competitive ability, and particularly against biogeographically new neighbors, but one endophyte increased the competitive ability of C. stoebe without increasing its size, suggesting mechanisms unrelated to increased growth. We tested only a fraction of the different endophytic fungi that have been found in C. stoebe, only scratching the surface of understanding their indirect effects. However, our results are the first to demonstrate such effects of a fungal endophyte infecting an invasive forb, and one of the few to show that endophyte effects on competition do not have to be mediated through herbivory.     

Simulated green turtle grazing affects nutrient composition of the seagrass <Emphasis Type="Italic">Thalassia testudinum</Emphasis>

Before populations of green turtles (Chelonia mydas) were severely reduced by human overexploitation, the seagrass Thalassia testudinum was intensively grazed by green turtles in the Caribbean. To explore how nutrient composition of T. testudinum pastures responds to intense grazing pressure, we simulated green turtle grazing in 15 plots (each 3 m × 3 m) for 16 months in the central Bahamas. Comparisons of clipped plots with 15 adjacent control (unclipped) plots revealed that simulated grazing resulted in significantly higher energy, nitrogen, phosphorus, lignin, cutin, and condensed tannin content in blades in clipped plots, but sediment organic content was not affected. By continually re-cropping blades in grazing plots, turtles ingest young, actively growing blade tissue with higher energy, nitrogen, and phosphorus concentrations. Our 16-month clipping trial did not generate the expected decline in nutrient content in T. testudinum blades under intensive grazing. However, significant decreases in nitrogen and organic matter reserves in rhizomes, with declines apparent after 16 and 11 months, respectively, indicate that nutrient content of blades and/or blade productivity may decline under continued clipping.     

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.     

Vegetation pattern related tc grazing pressure in alpine meadows of Nanda Devi Biosphere Reserve

The present study aims to analyze the interaction of prevailing biotic pressure on plant species diversity in Nanda Devi Biosphere Reserve (NDBR) which lies in northern part of Uttaranchal hills between 79 degrees 40'E to 80 degrees 05'E longitude and 30 degrees 17' N to 30 degrees 41'N latitude and covers an area of 2236.7 km2. A total of 75 species has been found which included the herbaceous plants viz., grasses, sedges and forbs. Generally, the plants have a short life span of 3-4 months. However, few species persist throughout the growth period i.e. May-October. Phytosociological study performed in plots of varying slope and grazing pressure intensity revealed that the dominant grasses were Danthonia cachemyriana and Poa annua and dominant forbs were Trachydium roylei and Geum elatum in all the plots. Grasses were abundant on west facing slopes while forbs preferred the even topography of east facing meadows. The grasses and sedges together had optimum density during July and August. In general, short lived species exhibited more diversity for one or two months whilst the long lived species exhibited optimum diversity althrough the snow free period. The species diversity is maximum (100%) in moderately grazed bughiyals i.e. Pacchu and minimum in intensively grazed bughiyals i.e. Martoli. The species distribution among the plots was 60-90% contagious and 11.2-38.0% randomirrespective of grazing pressure, thus highlighting the significance of grazing pressure in management of alpine meadows.     

Are soil mite assemblages structured by the identity of native and invasive alien grasses?

Associations between plants and animals in aboveground communities are often predictable and specific. This has been exploited for the purposes of estimating the diversity of animal species based on the diversity of plant species. The introduction of invasive alien plants into an ecosystem can result in dramatic changes in both the native plant and animal assemblages. Few data exist at the species level to determine whether belowground animal assemblages share the same degree of association to plants. The hypotheses that soil mites (Acari) form assemblages specifically associated with different native grass species in an unmanipulated natural ecosystem and that invasive alien grasses will impact soil mite assemblage composition in this setting were tested. Soil mites sampled beneath five native and two invasive alien species of grasses at the Konza Prairie Biological Station, Kansas, USA, were similarly abundant, species rich, diverse, and taxonomically distinct. No mite species had affinities for a specific grass species. There was no evidence from analysis of similarity, canonical correspondence analysis, or a nonparametric assemblage analysis that the assemblage composition of soil mites was specific to grass species. Results suggest that soil mite assemblages were more related to characteristics of the plant assemblage as a whole or prevailing soil conditions. The most recent invasive alien grass did not support a successionally younger mite fauna, based on the ratio of mesostigmatid to oribatid mites, and neither of the two invasive grasses influenced mite assemblage structure, possibly because they had not yet substantially altered the soil environment. Our results suggest that extrapolations of soil mite diversity based on assumptions of plant specificity would be invalid.     

刈割对香港草地干物质生产的影响

研究了刈割频率对香港草地于物质生产的影响。结果表明，刈割使干物质生产量下降，刈割频率越高生产量下降越大。禾本科植物受刈割影响比木本和非禾本科草本植物大，且停止刚割一年后生物量仍低于对照点。认为常年刈割是华南草地生产力下降的重要原因之一。     

Soil feedback of exotic savanna grass relates to pathogen absence and mycorrhizal selectivity

Enemy release of exotic plants from soil pathogens has been tested by examining plant-soil feedback effects in repetitive growth cycles. However, positive soil feedback may also be due to enhanced benefit from the local arbuscular mycorrhizal fungi (AMF). Few studies actually have tested pathogen effects, and none of them did so in arid savannas. In the Kalahari savanna in Botswana, we compared the soil feedback of the exotic grass Cenchrus biflorus with that of two dominant native grasses, Eragrostis lehmanniana and Aristida meridionalis. The exotic grass had neutral to positive soil feedback, whereas both native grasses showed neutral to negative feedback effects. Isolation and testing of root-inhabiting fungi of E. lehmanniana yielded two host-specific pathogens that did not influence the exotic C. biflorus or the other native grass, A. meridionalis. None of the grasses was affected by the fungi that were isolated from the roots of the exotic C. biflorus. We isolated and compared the AMF community of the native and exotic grasses by polymerase chain reaction-denaturing gradient gel elecrophoresis (PCR-DGGE), targeting AMF 18S rRNA. We used roots from monospecific field stands and from plants grown in pots with mixtures of soils from the monospecific field stands. Three-quarters of the root samples of the exotic grass had two nearly identical sequences, showing 99% similarity with Glomus versiforme. The two native grasses were also associated with distinct bands, but each of these bands occurred in only a fraction of the root samples. The native grasses contained a higher diversity of AMF bands than the exotic grass. Canonical correspondence analyses of the AMF band patterns revealed almost as much difference between the native and exotic grasses as between the native grasses. In conclusion, our results support the hypothesis that release from soil-borne enemies may facilitate local abundance of exotic plants, and we provide the first evidence that these processes may occur in arid savanna ecosystems. Pathogenicity tests implicated the involvement of soil pathogens in the soil feedback responses, and further studies should reveal the functional consequences of the observed high infection with a low diversity of AMF in the roots of exotic plants.     

Effects of Long-Term Ungulate Exclusion and Recent Alien Species Control on the Preservation and Restoration of a Hawaiian Tropical Dry Forest

Abstract: Although the destruction of tropical rain forests receives much attention, tropical dry forests are in general far more threatened and endangered. Eliminating grazing ungulates is often considered a key first step toward protecting these ecosystems, but few studies have investigated the long-term effects of this technique. We examined the effects of ungulate exclusion from a 2.3-ha native dry-forest preserve on the island of Hawaii by comparing its present flora to the flora of an adjacent area subjected to continuous grazing since the preserve was fenced over 40 years ago. Relative to this adjacent area, the fenced preserve contained a more diverse flora with substantially greater coverage of native overstory and understory species. Until recently, however, regeneration of native canopy trees within the preserve appears to have been thwarted by a dominant herbaceous cover of alien fountain grass (   Pennisetum setaceum ) and predation by alien rodent species. Our results indicate that although ungulate exclusion may be a necessary and critical first step, it is not sufficient to adequately preserve and maintain Hawaii's remaining tropical dry forest remnants. Our recent efforts to control the dominant alien species within the fenced preserve suggest that this practice may facilitate both the regeneration of native species and the colonization and potential invasion of new alien plants. Comparisons of seedlings of the dominant native canopy tree Diospyros sandwicensis growing in sites both dominated by and free of fountain grass suggested that fountain grass inhibits Diospyros seedling growth and photosynthesis but may increase survival if seedlings are protected from ungulates.     

Plant biomass allocation strategies of the dominant species in an alpine meadow of northwestern Sichuan,China北大核心CSCD

Plant biomass partitioning is an important driver of whole-plant net carbon gain, as biomass allocation could directly affect plant's future growth and reproduction. Alpine meadow in the northwestern Sichuan was impressed by the abundant community structure and species diversity. This study on biomass allocation pattern of different functional types and lifeforms might help understand plant life-history strategy of alpine meadow plants. We investigated 72 dominant herbaceous species for their compartments, biomass, and morphological traits during 2012-2014. These plants were sampled from natural grassland, disturbed grassland, and wintergreen grassland; they belonged to three functional types (grass, sedge, and forb) and two lifeforms (annual and perennial). The scaling relationships between functional traits of these plants were analyzed using Model type II regression method to estimate the parameters of the allometric equations. (1) Biomass allocation proportion of components significantly differed among grasses, sedges, and forbs owing to phylogeny: grasses had the highest stem biomass percentage, sedges had higher root biomass percentage, and forbs had higher leaf biomass percentage, but the scaling relationships were not significantly different, and isometric scaling was noted between biomass components for the three functional types. (2) Moreover, plant lifeforms affected the biomass allocation proportion of components, owing to the shorter or longer turnover rate and investment strategy between annual and perennial species. Annuals allocated more biomass to the stem and reproduction organs, but perennials invested more biomass to the leaves and roots. (3) In addition, plants from different grassland types differed in both biomass and morphology traits. Moreover, forbs from natural grassland and wintergreen grassland had higher leaf and reproductive biomass, but those from disturbed grasslands had higher stem biomass. Our results suggest that the functional type and lifeform decide the inherent scaling relationships between components of plants, but anthropogenic disturbance significantly impacted the quantity of component biomass. This study has important theoretical and practical significance to understand the response of alpine plants to climate change and anthropogenic disturbance as well as to help in the scientific management of alpine meadow. © 2018 Science Press. All rights reserved.     

异质环境下入侵植物薇甘菊的适应性与繁殖特性

薇甘菊（Mikaniamicrantha H.B.Kunth）为世界上最具有入侵性和危害性的外来入侵物种之一,对其控制与管理已成为长期以来世界性难题。了解入侵植物薇甘菊在异质环境下的适应性与繁殖特性对安全有效预警、监测和防治该入侵物种具有重要意义。本文于薇甘菊盛花期,根据薇甘菊入侵生境的光照条件、群落结构和生长方式差异,在薇甘菊常见的4种入侵生境（林地、荒地、农田和和河边）选取10个样地,调查研究了不同生境条件下薇甘菊的开花结实和繁殖分配,并运用植物叶片功能性状的研究方法研究了不同生境条件下薇甘菊的比叶面积（SLA）、叶干物质比例（LMF）和茎干物质比例（SMF）的3种叶片功能性状。研究结果表明,生境条件对薇甘菊的开花结实和繁殖分配具有显著影响。其中抛荒农田（样地Ⅷ）和河流边缘（样地Ⅸ和Ⅹ）薇甘菊的总花数、种子量、生殖枝茎生物量分配和花生物量分配均显著大于其他生境,显示在有利于其生长的条件下,薇甘菊种群倾向于有性繁殖;而在不利于薇甘菊生长的低光照林地（样地Ⅰ）和种间竞争强的农田红薯地（样地Ⅵ）生境条件下,薇甘菊的总花数和种子量明显小于其他生境,但营养枝叶生物量分配显著大于其他样地,而茎生物量分配值则处于中间值,表明在不利于其生长的生境条件下,薇甘菊通过提高营养枝茎的生物量分配和叶的生物量分配来适应,其种群则更倾向于克隆繁殖。通过分析不同生境条件下薇甘菊的叶片功能性状,结果表明,在不利于薇甘菊生长的低光照林地（样地Ⅰ）和种间竞争强的农田红薯地（样地Ⅵ）生境条件下,薇甘菊的叶面积和茎干物质比例（SMF）显著小于其他样地,且彼此差异不显著;但比叶面积（SLA）和叶干物质比例（LMF）则显著大于其他样地,显示薇甘?     

Interplay between Senecio jacobaea and plant, soil, and aboveground insect community composition

To elucidate the factors that affect the performance of plants in their natural environment, it is essential to study interactions with other neighboring plants, as well as with above- and belowground higher trophic organisms. We used a long-term field experiment to study how local plant community diversity influenced colonization by the biennial composite Senecio jacobaea in its native range in The Netherlands in Europe. We tested the effect of sowing later-succession plant species (0, 4, or 15 species) on plant succession and S. jacobaea performance. Over a period of eight years, the percent cover of S. jacobaea was relatively low in communities sown with 15 or 4 later-succession plant species compared to plots that were not sown, but that were colonized naturally. However, after four years of high abundance, the density of S. jacobaea in unsown plots started to decline, and the size of the individual plants was smaller than in the plots sown with 15 or 4 plant species. In the unsown plots, densities of aboveground leaf-mining, flower-feeding, and stem-boring insects on S. jacobaea plants were lower than on plants in sown plots, and there was a strong positive relationship between plant size and levels of herbivory. In a greenhouse experiment, we grew S. jacobaea in sterilized soil inoculated with soil from the different sowing treatments of the field experiment. Biomass production was lower when S. jacobaea test plants were grown in soil from the unsown plots than in soil from the sown plots (4 or 15 species). Molecular analysis of the fungal and bacterial communities revealed that the composition of fungal communities in unsown plots differed significantly from those in sown plots, suggesting that soil fungi could have been involved in the relative growth reduction of S. jacobaea in the greenhouse bioassay. Our results show that, in its native habitat, the abundance of S. jacobaea depends on the initial composition of the plant community and that, on a scale of almost a decade, its interactions with plant and soil communities and aboveground invertebrates may influence the dynamics of this colonizing species.     

Gastropod herbivory in response to elevated CO2 and N addition impacts plant community composition

In this study, the influence of elevated carbon dioxide (CO2) and nitrogen (N) deposition on gastropod herbivory was investigated for six annual species in a California annual grassland community. These experimentally simulated global changes increased availability of important resources for plant growth, leading to the hypothesis that species with the most positive growth and foliar nutrient responses would experience the greatest increase in herbivory. Counter to the expectations, shifts in tissue N and growth rates caused by N deposition did not predict shifts in herbivore consumption rates. N deposition increased seedling N concentrations and growth rates but did not increase herbivore consumption overall, or for any individual species. Elevated CO2 did not influence growth rates nor have a statistically significant influence on seedling N concentrations. Elevated CO2 at ambient N levels caused a decline in the number of seedlings consumed, but the interaction between CO2 and N addition differed among species. The results of this study indicate that shifting patterns of herbivory will likely influence species composition as environmental conditions change in the future; however, a simple trade-off between shifting growth rates and palatability is not evident.     

Spatial and temporal patterns of seed dispersal: an important determinant of grassland invasion.

We measured spatial and temporal patterns of seed dispersal and seedling recruitment for 58 species in a grassland community to test whether seed dispersal could predict patterns of invasion after disturbance. For the 12 most abundant grasses, recruitment of native species was dependent on the propagule supply of both native and exotic species. Variability in seed rain on small spatial (1-10 m) and temporal (within season) scales led to qualitative differences in the outcome of disturbance colonization such that native species dominated disturbances when exotic seed supply was low but failed to establish when exotic seed supply was high. Local dispersal and spatial heterogeneity in species composition promoted coexistence of native and exotic species by creating refuges from high exotic seed supply within native dominated patches. Despite this, copious exotic seed production strongly limited recruitment of native species in exotic dominated patches. Most grasslands in California are presently dominated by exotic species, suggesting that competition at the seedling stage is a major barrier to native species restoration.     

Population-level compensation impedes biological control of an invasive forb and indirect release of a native grass

The intentional introduction of specialist insect herbivores for biological control of exotic weeds provides ideal but understudied systems for evaluating important ecological concepts related to top-down control, plant compensatory responses, indirect effects, and the influence of environmental context on these processes. Centaurea stoebe (spotted knapweed) is a notorious rangeland weed that exhibited regional declines in the early 2000s, attributed to drought by some and to successful biocontrol by others. We initiated an experiment to quantify the effects of the biocontrol agent, Cyphocleonus achates, on Ce. stoebe and its interaction with a dominant native grass competitor, Pseudoroegneria spicata, under contrasting precipitation conditions. Plots containing monocultures of each plant species or equal mixtures of the two received factorial combinations of Cy. achates herbivory (exclusion or addition) and precipitation (May-June drought or "normal," defined by the 50-year average) for three years. Cy. achates herbivory reduced survival of adult Ce. stoebe plants by 9% overall, but this effect was stronger under normal precipitation compared to drought conditions, and stronger in mixed-species plots compared to monocultures. Herbivory had no effect on Ce. stoebe per capita seed production or on recruitment of seedlings or juveniles. In normal-precipitation plots of mixed composition, greater adult mortality due to Cy. achates herbivory resulted in increased recruitment of new adult Ce. stoebe. Due to this compensatory response to adult mortality, final Ce. stoebe densities did not differ between herbivory treatments regardless of context. Experimental drought reduced adult Ce. stoebe survival in mixed-species plots but did not impede recruitment of new adults or reduce final Ce. stoebe densities, perhaps due to the limited duration of the treatment. Ce. stoebe strongly depressed P. spicata reproduction and recruitment, but these impacts were not substantively alleviated by herbivory on Ce. stoebe. Population-level compensation by dominant plants may be an important factor inhibiting top-down effects in herbivore-driven and predator-driven cascades.