Grassland invader responses to realistic changes in native species richness.

The importance of species richness for repelling exotic plant invasions varies from ecosystem to ecosystem. Thus, in order to prioritize conservation objectives, it is critical to identify those ecosystems where decreasing richness will most greatly magnify invasion risks. Our goal was to determine if invasion risks greatly increase in response to common reductions in grassland species richness. We imposed treatments that mimic management-induced reductions in grassland species richness (i.e., removal of shallow- and/or deep-rooted forbs and/or grasses and/or cryptogam layers). Then we introduced and monitored the performance of a notorious invasive species (i.e., Centaurea maculosa). We found that, on a per-gram-of-biomass basis, each resident plant group similarly suppressed invader growth. Hence, with respect to preventing C. maculosa invasions, maintaining overall productivity is probably more important than maintaining the productivity of particular plant groups or species. But at the sites we studied, all plant groups may be needed to maintain overall productivity because removing forbs decreased overall productivity in two of three years. Alternatively, removing forbs increased productivity in another year, and this led us to posit that removing forbs may inflate the temporal productivity variance as opposed to greatly affecting time-averaged productivity. In either case, overall productivity responses to single plant group removals were inconsistent and fairly modest, and only when all plant groups were removed did C. maculosa growth increase substantially over a no-removal treatment. As such, it seems that intense disturbances (e.g., prolonged drought, overgrazing) that deplete multiple plant groups may often be a prerequisite for C. maculosa invasion.     

An invader differentially affects leaf physiology of two natives across a gradient in diversity

Little is known about how exotics influence the ecophysiology of co-occurring native plants or how invader impact on plant physiology may be mediated by community diversity or resource levels. We measured the effect of the widespread invasive forb spotted knapweed (Centaurea maculosa) on leaf traits (leaf dry matter content, specific leaf area, leaf nitrogen percentage, leaf C:N ratios, and delta13C as a proxy for water use efficiency) of two co-occurring native perennial grassland species, Monarda fistulosa (bee balm) and Koeleria macrantha (Junegrass). The impact of spotted knapweed was assessed across plots that varied in functional diversity and that either experienced ambient rainfall or received supplemental water. Impact was determined by comparing leaf traits between identical knapweed-invaded and noninvaded assemblages. Virtually all M. fistulosa leaf traits were affected by spotted knapweed. Knapweed impact, however, did not scale with its abundance; the impact of knapweed on M. fistulosa was similar across heavily invaded low-diversity assemblages and lightly invaded high-diversity assemblages. In uninvaded assemblages, M. fistulosa delta13C, leaf nitrogen, and C:N ratios were unaffected by native functional group richness, whereas leaf dry matter content significantly increased and specific leaf area significantly decreased across the diversity gradient. The effects of spotted knapweed on K. macrantha were weak; instead native functional group richness strongly affected K. macrantha leaf C:N ratio, delta13C, and specific leaf area, but not leaf dry matter content. Leaf traits for both species changed in response to spotted knapweed or functional richness, and in a manner that may promote slower biomass accumulation and efficient conservation of resources. Taken together, our results show that an invader can alter native plant physiology, but that these effects are not a simple function of how many invaders exist in the community.     

Indices of Grassland Biodiversity in the Chihuahuan Desert Ecoregion Derived from Remote Sensing

Abstract: We used a relatively simple and direct remote-sensing approach to determine biodiversity values in arid ecosystems and thus identify potential conservation sites. We developed indices based on regression models between grass, shrub, litter, exposed-soil groundcover components, and Landsat thematic mapper satellite imagery reflectance values over a reference site in the northern Chihuahuan Desert in New Mexico. This site supports low-disturbance desert grasslands that have been excluded from livestock grazing for 55 years and moderate-disturbance grasslands that have been under a continuous grazing regime for over 100 years. Greater richness and abundance of noninvasive and nonruderal plant species were associated with the low-disturbance grasslands that had lower shrub abundance, increased litter and grass cover, and lower exposed soil. Using the thematic mapper indices, we computed an additive grassland biodiversity index such that, as exposed soil and shrub values go down, litter and grass values go up, as does the biodiversity index. When the biodiversity index was applied to the reference-site landscape, grasslands previously identified for their high conservation value were detected. As a further test, we applied the indices to a site in Chihuahua, Mexico, that supports similar grasslands but for which there are few other data on condition and conservation values. The soil, grass, and shrub indices were moderately effective in describing the range of variation at the Mexico site, but the litter equation was not. Still, higher biodiversity value in terms of nonruderal plant diversity tended to correspond to higher grass cover and lower soil exposure and a higher overall biodiversity index. Some localized calibration with geologic substrate may be required along with an assessment of the temporal constraints, but generally the index shows promise for quickly and efficiently detecting desert grasslands of high biodiversity conservation value.     

Postfire Seeding for Erosion Control: Effectiveness and Impacts on Native Plant Communities

Abstract:  Large, high-severity wildfires remove vegetation cover and expose mineral soil, ofen causing erosion and runoff during postfire rain events to increase dramatically. Land-management agencies in the United States are required to assess site conditions after wildfire and, where necessary, implement emergency watershed rehabilitation measures to help stabilize soil; control movement of water, sediment, and debris; prevent permanent impairment of ecosystem structure and function; and mitigate significant threats to human health, safety, life, property, or downstream values. One of the most common postfire treatments is broadcast seeding of grasses, usually from aircraft. Non-native annual or perennial grasses typically are used to provide quick, temporary ground cover to hold soil in place until native plants are reestablished. Critics argue that seeded grasses compete with native vegetation and do not effectively reduce erosion. Few data exist on the effectiveness of erosion control; less than half of the studies I reviewed showed reduced sediment movement with seeding. In all vegetation types, successful growth of seeded grasses—enough to affect erosion—appears to displace native or naturalized species, including shrub and tree seedlings. Due to the competitiveness of seeded grasses, they are used to attempt suppression of noxious weeds in some postfire seeding operations. In burned sagebrush range, postfire seeding is frequently used to replace non-native cheatgrass (  Bromus tectorum ) with native or introduced bunchgrasses, with at least short-term success. In recent years, native species and sterile cereal grains have increasingly been used for seeding. Use of aerially applied straw mulch has increased as well, with the risk of weed introduction from contaminated bales. More research on the effectiveness and ecosystem impacts of these alternatives is needed.     

Plant winners and losers during grassland N-eutrophication differ in biomass allocation and mycorrhizas

Human activities release tremendous amounts of nitrogenous compounds into the atmosphere. Wet and dry deposition distributes this airborne nitrogen (N) on otherwise pristine ecosystems. This eutrophication process significantly alters the species composition of native grasslands; generally a few nitrophilic plant species become dominant while many other species disappear. The functional equilibrium model predicts that, compared to species that decline in response to N enrichment, nitrophilic grass species should respond to N enrichment with greater biomass allocation aboveground and reduced allocation to roots and mycorrhizas. The mycorrhizal feedback hypothesis states that the composition of mycorrhizal fungal communities may influence the composition of plant communities, and it predicts that N enrichment may generate reciprocal shifts in the species composition of mycorrhizal fungi and plants. We tested these hypotheses with experiments that compared biomass allocation and mycorrhizal function of four grass ecotypes (three species), two that gained and two that lost biomass and cover in response to long-term N enrichment experiments at Cedar Creek and Konza Long-Term Ecological Research grasslands. Local grass ecotypes were grown in soil from their respective sites and inoculated with whole-soil inoculum collected from either fertilized (FERT) or unfertilized (UNFERT) plots. Our results strongly support the functional equilibrium model. In both grassland systems the nitrophilic grass species grew taller, allocated more biomass to shoots than to roots, and formed fewer mycorrhizas compared to the grass species that it replaced. Our results did not fully support the hypothesis that N-induced changes in the mycorrhizal fungal community were drivers of the plant community shifts that accompany N eutrophication. The FERT and UNFERT soil inoculum influenced the growth of the grasses differently, but this varied with site and grass ecotype in both expected and unexpected ways suggesting that ambient soil fertility or other factors may be interacting with mycorrhizal feedbacks.     

Effects of initial irrigation,seeding date and directlyhauled topsoil on mined land community composition

This study was implemented to determine the effects of one, two, or three months of establishment irrigation, after spring or summer seeding on the long-term composition of mined land plant associations. While the treatment plots were seeded with the same diverse mixture of native species, floristic differences among treatment combinations were apparent after five growing seasons. Summer seeding enhanced the development of seeded warm-season grasses by retarding the initial development of seeded cool-season grasses and by limiting competitive effects of nonseeded species through seedbed tillage. Supplemental irrigation was substantially more important to the development of warm- than cool-season grasses. Seeded perennial forbs performed poorly on all treatments, presumably due to low seed viability, low germination, and/or competition from more water-responsive species. Despite drought conditions during plant establishment, supplemental irrigation was not critical to the ultimate development of nonseeded plant species when seedbed tillage occurred shortly after topsoil was applied. However, irrigation became more important to the long-term development of these species when tillage was delayed. Most forbs encountered were perennial natives that had volunteered from the direct-haul topsoil.Atriplex canescens was the only successful seeded shrub and performed particularly well on the summer seeded plot that received one month of initial irrigation.Authors are research plant ccologist and director of research. This contribution submitted as Montana Agricultural Experiment Station Journal Series No. J-1560. This study was funded entirely by the Western Energy Company, Butte, Montana. The authors express their appreciation to Bill Schwarzkoph and Joe Cocnenberg of Western Energy Company, and former Reclamation Research Unit members Ed DePuit, Chet Skilbred, and Steve Young for their participation in the early stages of this study.     

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.     

Weed-biocontrol insects reduce native-plant recruitment through second-order apparent competition

Small-mammal seed predation is an important force structuring native-plant communities that may also influence exotic-plant invasions. In the intermountain West, deer mice (Peromyscus maniculatus) are prominent predators of native-plant seeds, but they avoid consuming seeds of certain widespread invasives like spotted knapweed (Centaurea maculosa). These mice also consume the biological-control insects Urophora spp. introduced to control C. maculosa, and this food resource substantially increases deer mouse populations. Thus, mice may play an important role in the invasion and management of C. maculosa through food-web interactions. We examined deer mouse seed predation and its effects on seedling emergence and establishment of a dominant native grass, Pseudoroegneria spicata, and forb, Balsamorhiza sagittata, in C. maculosa-invaded grasslands that were treated with herbicide to suppress C. maculosa or left untreated as controls. Deer mice readily took seeds of both native plants but removed 2-20 times more of the larger B. sagittata seeds than the smaller P. spicata seeds. Seed predation reduced emergence and establishment of both species but had greater impacts on B. sagittata. The intensity of seed predation corresponded with annual and seasonal changes in deer mouse abundance, suggesting that abundance largely determined mouse impacts on native-plant seeds. Accordingly, herbicide treatments that reduced mouse abundance by suppressing C. maculosa and its associated biocontrol food subsidies to mice also reduced seed predation and decreased the impact of deer mice on B. sagittata establishment. These results provide evidence that Urophora biocontrol agents may exacerbate the negative effects of C. maculosa on native plants through a form of second-order apparent competition-a biocontrol indirect effect that has not been previously documented. Herbicide suppressed C. maculosa and Urophora, reducing mouse populations and moderating seed predation on native plants, but the herbicide's direct negative effects on native forb seedlings overwhelmed the indirect positive effect of reducing deer mouse seed predation. By manipulating this four-level food chain, we illustrate that host-specific biological control agents may impact nontarget plant species through food-web interactions, and herbicides may influence management outcomes through indirect trophic interactions in addition to their direct effects on plants.     

The invasive grass Agropyron cristatum doubles belowground productivity but not soil carbon

Root dynamics are among the largest knowledge gaps in determining how terrestrial carbon (C) cycles will respond to environmental change. Increases in productivity accompanying plant invasions and introductions could increase ecosystem C storage, but belowground changes are unknown, even though roots may account for 50-90% of production in temperate ecosystems. We examined whether the introduction of a widespread invasive grass with relatively high shoot production also increased belowground productivity and soil C storage, using a multiyear rhizotron study in 50-year-old stands dominated either by the invasive C3 grass Agropyron cristatum or by largely C4 native grasses. Relative to native vegetation, stands dominated by the invader had doubled root productivity. Soil carbon isotope values showed that the invader had made detectable contributions to soil C. Soil C content, however, was not significantly different between invader-dominated stands (0.42 mg C/g soil) and native vegetation (0.45 mg C/g soil). The discrepancy between enhanced production and lack of soil C changes was attributable to differences in root traits between invader-dominated stands and native vegetation. Relative to native vegetation, roots beneath the invader had 59% more young white tissue, with 80% higher mortality and 19% lower C:N ratios (all P < 0.05). Such patterns have previously been reported for aboveground tissues of invaders, and we show that they are also found belowground. If these root traits occur in other invasive species, then the global phenomenon of increased productivity following biological invasion may not increase soil C storage.     

Forest succession suppressed by an introduced plant-fungal symbiosis

Microbial symbionts can affect plant nutrition, defensive chemistry, and biodiversity. Here we test the hypothesis that symbionts alter the speed and direction of plant succession in communities that are shifting from grasslands to forests. A widespread C3 grass introduced to the United States, Lolium arundinaceum (tall fescue), hosts a fungal endophyte that is toxic to herbivores. In replicated experimental grasslands, the presence of the endophyte in tall fescue reduced tree abundance and size, altered tree composition, and slowed plant species turnover. In addition, consumption of tree seedlings by voles (Microtus spp.) was 65% higher in plots with the endophyte at the one grassland site where these data were collected. Despite its negligible contribution to community biomass, a microbial symbiont suppressed tree establishment, posing an important constraint on the natural transition from grasslands to forests.     

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.     

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.     

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.     

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.     

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.     

Long-term dynamics of biotic and abiotic resistance to exotic species invasion in restored vernal pool plant communities

Invasion of native ecosystems by exotic species can seriously threaten native biodiversity, alter ecosystem function, and inhibit conservation. Moreover, restoration of native plant communities is often impeded by competition from exotic species. Exotic species invasion may be limited by unfavorable abiotic conditions and by competition with native species, but the relative importance of biotic and abiotic factors remains controversial and may vary during the invasion process. We used a long-term experiment involving restored vernal pool plant communities to characterize the temporal dynamics of exotic species invasion, and to evaluate the relative support for biotic and abiotic factors affecting invasion resistance. Experimental pools (n=256) were divided among controls and several seeding treatments. In most treatments, native vernal pool species were initially more abundant than exotic species, and pools that initially received more native seeds exhibited lower frequencies of exotic species over time. However, even densely seeded pools were eventually dominated by exotic species, following extreme climatic events that reduced both native and exotic plant densities across the study site. By the sixth year of the experiment, most pools supported more exotics than native vernal pool species, regardless of seeding treatment or pool depth. Although deeper pools were less invaded by exotic species, two exotics (Hordeum marinum and Lolium multiflorum) were able to colonize deeper pools as soon as the cover of native species was reduced by climatic extremes. Based on an information-theoretic analysis, the best model of invasion resistance included a nonlinear effect of seeding treatment and both linear and nonlinear effects of pool depth. Pool depth received more support as a predictor of invasion resistance, but seeding intensity was also strongly supported in multivariate models of invasion, and was the best predictor of resistance to invasion by H. marinum and L. multilorum. We conclude that extreme climatic events can facilitate exotic species invasions by both reducing abiotic constraints and weakening biotic resistance to invasion.     

Bromus tectorum invasion alters nitrogen dynamics in an undisturbed arid grassland ecosystem

The nonnative annual grass Bromus tectorum has successfully replaced native vegetation in many arid and semiarid ecosystems. Initial introductions accompanied grazing and agriculture, making it difficult to separate the effects of invasion from physical disturbance. This study examined N dynamics in two recently invaded, undisturbed vegetation associations (C3 and C4). The response of these communities was compared to an invaded/ disturbed grassland. The invaded/disturbed communities had higher surface NH4+ input in spring, whereas there were no differences for surface input of NO3-. Soil inorganic N was dominated by NH4+, but invaded sites had greater subsurface soil NO3-. Invaded sites had greater total soil N at the surface four years post-invasion in undisturbed communities, but total N was lower in the invaded/disturbed communities. Soil delta15N increased with depth in the noninvaded and recently invaded communities, whereas the invaded/disturbed communities exhibited the opposite pattern. Enriched foliar delta15N values suggest that Bromus assimilated subsurface NO3-, whereas the native grasses were restricted to surface N. A Rayleigh distillation model accurately described decomposition patterns in the noninvaded communities where soil N loss is accompanied by increasing soil delta15N; however, the invaded/ disturbed communities exhibited the opposite pattern, suggesting redistribution of N within the soil profile. This study suggests that invasion has altered the mechanisms driving nitrogen dynamics. Bromus litter decomposition and soil NO3- concentrations were greater in the invaded communities during periods of ample precipitation, and NO3- leached from the surface litter, where it was assimilated by Bromus. The primary source of N input in these communities is a biological soil crust that is removed with disturbance, and the lack of N input by the biological soil crust did not balance N loss, resulting in reduced total N in the invaded/disturbed communities. Bromus produced a positive feedback loop by leaching NO3- from decomposing Bromus litter to subsurface soil layers, accessing that deepsoil N pool with deep roots and returning that N to the surface as biomass and subsequent litter. Lack of new inputs combined with continued loss will result in lower total soil N, evidenced by the lower total soil N in the invaded/disturbed communities.     

Mitigating exotic impacts: restoring deer mouse populations elevated by an exotic food subsidy.

The threat posed by exotic organisms to native systems has led to extensive research on exotic invaders, yet management of invasives has progressed relatively slowly. This is partly due to poor understanding of how exotic species management influences native organisms. To address this shortfall, we experimentally evaluated the efficacy of an invasives management tool for restoring native deer mouse (Peromyscus maniculatus) populations elevated by exotic species. The exotic insects, Urophora spp., were introduced in North America for biological control of the Eurasian invader, spotted knapweed (Centaurea maculosa), but instead of controlling C. maculosa, Urophora have become an important food resource that doubles P. maniculatus populations, with substantial indirect effects on other organisms. We hypothesized that herbicide suppression of Urophora's host plant would reduce the Urophora food resource and restore P. maniculatus populations to natural levels. Prior to treatment, mouse populations did not differ between controls and treatments, but following treatment, P. maniculatus were half as abundant where treatment reduced Urophora. Peromyscus maniculatus is insensitive to direct herbicide effects, and herbicide-induced habitat changes could not explain the P. maniculatus response. Treatment-induced reductions of the Urophora food resource offered the most parsimonious explanation for the mouse response: Multistate mark-recapture models indicated that P. maniculatus survival declined where Urophora were removed, and survival rates were more correlated with variation in population size than movement rates. Other demographic and reproductive parameters (sex ratios, reproductive status, pregnancy rates, and juvenile recruitment) were unaffected by treatment. These results suggest the Urophora biocontrol elevated P. maniculatus survival, and the herbicide treatment restored mouse populations by removing the exotic food and reducing survival. This work illustrates the importance of mechanistic understandings of community and population ecology for improving invasive species management.     

Balancing Forest‐Regeneration Probabilities and Maintenance Costs in Dry Grasslands of High Conservation Priority

Abstract: Abandonment of agricultural land has resulted in forest regeneration in species‐rich dry grasslands across European mountain regions and threatens conservation efforts in this vegetation type. To support national conservation strategies, we used a site‐selection algorithm (MARXAN) to find optimum sets of floristic regions (reporting units) that contain grasslands of high conservation priority. We sought optimum sets that would accommodate 136 important dry‐grassland species and that would minimize forest regeneration and costs of management needed to forestall predicted forest regeneration. We did not consider other conservation elements of dry grasslands, such as animal species richness, cultural heritage, and changes due to climate change. Optimal sets that included 95–100% of the dry grassland species encompassed an average of 56–59 floristic regions (standard deviation, SD 5). This is about 15% of approximately 400 floristic regions that contain dry‐grassland sites and translates to 4800–5300 ha of dry grassland out of a total of approximately 23,000 ha for the entire study area. Projected costs to manage the grasslands in these optimum sets ranged from CHF (Swiss francs) 5.2 to 6.0 million/year. This is only 15–20% of the current total estimated cost of approximately CHF30–45 million/year required if all dry grasslands were to be protected. The grasslands of the optimal sets may be viewed as core sites in a national conservation strategy.