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.     

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.     

Evaluating Effects of Habitat Loss and Land-Use Continuity on Ant Species Richness in Seminatural Grassland Remnants

Abstract:  Seminatural grasslands in Europe are susceptible to habitat destruction and fragmentation that result in negative effects on biodiversity because of increased isolation and area effects on extinction rate. However, even small habitat patches of seminatural grasslands might be of value for conservation and restoration of species richness in a landscape with a long history of management, which has been argued to lead to high species richness. We tested whether ant communities have been negatively affected by habitat loss and increased isolation of seminatural grasslands during the twentieth century. We examined species richness and community composition in seminatural grasslands of different size in a mosaic landscape in Central Sweden. Grasslands managed continuously over centuries harbored species-rich and ecologically diverse ant communities. Grassland remnant size had no effect on ant species richness. Small grassland remnants did not harbor a nested subset of the ant species of larger habitats. Community composition of ants was mainly affected by habitat conditions. Our results suggest that the abandonment of traditional land use and the encroachment of trees, rather than the effects of fragmentation, are important for species composition in seminatural grasslands. Our results highlight the importance of considering land-use continuity and dispersal ability of the focal organisms when examining the effects of habitat loss and fragmentation on biodiversity. Landscape history should be considered in conservation programs focusing on effects of land-use change.     

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.     

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.     

Long-term population dynamics of seeded plants in invaded grasslands

In recent decades, dozens of studies have involved attempts to introduce native and desirable nonnative plant species into grasslands dominated by invasive weeds. The newly introduced plants have proved capable of establishing, but because they are rarely monitored for more than four years, it is unknown if they have a high likelihood of persisting and suppressing invaders for the long-term. Beyond invaded grasslands, this lack of long-term monitoring is a general problem plaguing efforts to reintroduce a range of taxa into a range of ecosystems. We introduced species from seed and then periodically measured plant abundances for nine years at one site and 15 years at a second site. To our knowledge, our 15-year data are the longest to date from a seeding experiment in invaded, never-cultivated grassland. At one site, three seeded grasses maintained high densities for three or more years, but then all or nearly all individuals died. At the second site, one grass performed similarly, but two other grasses proliferated and at least one greatly suppressed the dominant invader (Centaurea maculosa). In one study, our point estimate suggests that the seeded grass Thinopyrum intermedium reduced C. maculosa biomass by 93% 15 years after seeding. In some cases, data from three and fewer years after seeding falsely suggested that seeded species were capable of persisting within the invaded grassland. In other cases, data from as late as nine years after seeding falsely suggested seeded populations would not become large enough to suppress the invader. These results show that seeded species sometimes persist and suppress invaders for long periods, but short-term data cannot predict if, when, or where this will occur. Because short-term data are not predictive of long-term seeded species performances, additional long-term data are needed to identify effective practices, traits, and species for revegetating invaded grasslands.     

Mycorrhizal fungal identity and diversity relaxes plant-plant competition

There is a great interest in ecology in understanding the role of soil microbial diversity for plant productivity and coexistence. Recent research has shown increases in species richness of mutualistic soil fungi, the arbuscular mycorrhizal fungi (AMF), to be related to increases in aboveground productivity of plant communities. However, the impact of AMF richness on plant-plant interactions has not been determined. Moreover, it is unknown whether species-rich AMF communities can act as insurance to maintain productivity in a fluctuating environment (e.g., upon changing soil conditions). We tested the impact of four different AMF taxa and of AMF diversity (no AMF, single AMF taxa, and all four together) on competitive interactions between the legume Trifolium pratense and the grass Lolium multiflorum grown under two different soil conditions of low and high sand content. We hypothesized that more diverse mutualistic interactions (e.g., when four AMF taxa are present) can ease competitive effects between plants, increase plant growth, and maintain plant productivity across different soil environments. We used quantitative PCR to verify that AMF taxa inoculated at the beginning of the experiment were still present at the end. The presence of AMF reduced the competitive inequality between the two plant species by reducing the growth suppression of the legume by the grass. High AMF richness enhanced the combined biomass production of the two plant species and the yield of the legume, particularly in the more productive soil with low sand content. In the less productive (high sand content) soil, the single most effective AMF had an equally beneficial effect on plant productivity as the mixture of four AMF. Since contributions of single AMF to plant productivity varied between both soils, higher AMF richness would be required to maintain plant productivity in heterogeneous environments. Overall this work shows that AMF diversity promotes plant productivity and that AMF diversity can act as insurance to sustain plant productivity under changing environmental conditions.     

Predicting leaf physiology from simple plant and climate attributes: a global GLOPNET analysis.

Knowledge of leaf chemistry, physiology, and life span is essential for global vegetation modeling, but such data are scarce or lacking for some regions, especially in developing countries. Here we use data from 2021 species at 175 sites around the world from the GLOPNET compilation to show that key physiological traits that are difficult to measure (such as photosynthetic capacity) can be predicted from simple qualitative plant characteristics, climate information, easily measured ("soft") leaf traits, or all of these in combination. The qualitative plant functional type (PFT) attributes examined are phylogeny (angiosperm or gymnosperm), growth form (grass, herb, shrub, or tree), and leaf phenology (deciduous vs. evergreen). These three PFT attributes explain between one-third and two-thirds of the variation in each of five quantitative leaf ecophysiological traits: specific leaf area (SLA), leaf life span, mass-based net photosynthetic capacity (Amass), nitrogen content (N(mass)), and phosphorus content (P(mass)). Alternatively, the combination of four simple, widely available climate metrics (mean annual temperature, mean annual precipitation, mean vapor pressure deficit, and solar irradiance) explain only 5-20% of the variation in those same five leaf traits. Adding the climate metrics to the qualitative PFTs as independent factors in the model increases explanatory power by 3-11% for the five traits. If a single easily measured leaf trait (SLA) is also included in the model along with qualitative plant traits and climate metrics, an additional 5-25% of the variation in the other four other leaf traits is explained, with the models accounting for 62%, 65%, 66%, and 73% of global variation in N(mass), P(mass), A(mass), and leaf life span, respectively. Given the wide availability of the summary climate data and qualitative PFT data used in these analyses, they could be used to explain roughly half of global variation in the less accessible leaf traits (A(mass), leaf life span, N(mass), P(mass)); this can be augmented to two-thirds of all variation if climatic and PFT data are used in combination with the readily measured trait SLA. This shows encouraging possibilities of progress in developing general predictive equations for macro-ecology, global scaling, and global modeling.     

Decomposers (Lumbricidae, Collembola) affect plant performance in model grasslands of different diversity

Decomposer invertebrates influence soil structure and nutrient mineralization as well as the activity and composition of the microbial community in soil and therefore likely affect plant performance and plant competition. We established model grassland communities in a greenhouse to study the interrelationship between two different functional groups of decomposer invertebrates, Lumbricidae and Collembola, and their effect on plant performance and plant nitrogen uptake in a plant diversity gradient. Common plant species of Central European Arrhenatherion grasslands were transplanted into microcosms with numbers of plant species varying from one to eight and plant functional groups varying from one to four. Separate and combined treatments with earthworms and collembolans were set up. Microcosms contained 15N labeled litter to track N fluxes into plant shoots. Presence of decomposers strongly increased total plant and plant shoot biomass. Root biomass decreased in the presence of collembolans and even more in the presence of earthworms. However, it increased when both animal groups were present. Also, presence of decomposers increased total N concentration and 15N enrichment of grasses, legumes, and small herbs. Small herbs were at a maximum in the combined treatment with earthworms and collembolans. The impact of earthworms and collembolans on plant performance strongly varied with plant functional group identity and plant species diversity and was modified when both decomposers were present. Both decomposer groups generally increased aboveground plant productivity through effects on litter decomposition and nutrient mineralization leading to an increased plant nutrient acquisition. The non-uniform effects of earthworms and collembolans suggest that functional diversity of soil decomposer animals matters and that the interactions between soil animal functional groups affect the structure of plant communities.     

An individual-based analysis of the dynamics of two coexisting phytoplankton species in the mixed layer

In marine ecosystems ecological and environmental conditions continuously change, possibly supporting the wide range of phytoplankton species coexisting in aquatic environments. Phytoplankton communities are not homogeneously distributed in the water column due to the spatial and temporal variability of turbulent mixing and the concurrent biological response. In this paper an individual-based model (Lagrangian method) simulating the basic physiology of two coexisting phytoplankton species has been developed. The species, sharing the same availability of light and nutrient resource, are characterized by different photo-physiological parameters. The spatial and temporal evolution of turbulent mixing is simulated introducing vertical profiles of measured eddy diffusivity. Three case studies have been examined to analyze the role of environment–individual interactions in determining bloom conditions for both the selected species. The organisms experience recurrent fluctuations of light, temperature, and nutrient concentration gradients, due to the turbulent mixing in the water column, which have significant effects on the growth of the phytoplankton species. In all the numerical experiments, the temporal and spatial variability of different forcings do not support the prevalence of one species over the other over the time scale typical of a phytoplankton bloom.A well mixed water column favours the growth of both the populations while a variable mixing regime limits their growth reducing the photophysiological differences between the species.     

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.     

Simulation of interrelations of the Everglades' marsh,peat, fire,water and phosphorus

Some of the principal controlling factors affecting the Everglades marsh system were combined in a simple model including growth of grass, water levels, rain, transpiration, peat deposition, fire and phosphorus and controlled inflow of water containing nutrients. Using data from published sources, coefficients were estimated and the model was simulated for several regimes, for varying concentrations of nutrient in the inflows and for varying access to fire. The resulting graphs resemble patterns reported from the Everglades, with some regimes producing regular repeating patterns and frequent small fires, whereas other regimes produced erratic and widely fluctuating patterns of vegetation, flood and fire. High phosphorus increased water loss by affecting plant transpiration: oscillations of vegetation and fire caused nutrient oscillations. Inflow of low nutrient water decreased nutrient levels by binding nutrients in plant masses. Continuous high water levels developed large accumulations of vegetation and peat, binding the nutrients, making larger fires when water levels were lowered. If this model is pertinent, a regular period of variation of water inflow and limited nutrients may be means for management of marshes for long range stability.     

Decline in metallophytes in tertiary polluted floodplain grasslands in the Netherlands: experimental evidence for metal and nutritional changes in soil as driver factors

Metallophyte vegetation along the River Geul has been almost completely replaced by grasses during the last decades. Field investigations indicated that this was accompanied by higher alkalinity and phosphate availability in the soil, related to the closure of the metal industry and intensification of agricultural practices. An experiment with a full factorial design for phosphate and zinc availability indicated that the metallophytes Silene vulgaris and Thlaspi caerulescens did not grow on zinc-poor soils, irrespective of phosphate availability. The grass Holcus lanatus performed well on phosphate-rich soil, irrespective of zinc availability. An experiment with zinc-poor and zinc-rich floodplain soils confirmed the high zinc demand of the metallophytes T. caerulescens and Armeria maritima and the zinc independence of H. lanatus. A third experiment indicated that a reduced zinc availability due to liming affected only the metallophyte T. caerulescens; it had no effect on the growth of the grass Festuca rubra. This means that increasing alkalinity leads to a decrease in zinc availability, limiting the growth of at least some metallophyte species. An increase in phosphate availability stimulates growth in more competitive fast-growing grasses under zinc-rich as well as zinc-poor conditions.     

Biophysical feedback mediates effects of invasive grasses on coastal dune shape

Vegetation at the aquatic-terrestrial interface can alter landscape features through its growth and interactions with sediment and fluids. Even similar species may impart different effects due to variation in their interactions and feedbacks with the environment. Consequently, replacement of one engineering species by another can cause significant change in the physical environment. Here we investigate the species-specific ecological mechanisms influencing the geomorphology of U.S. Pacific Northwest coastal dunes. Over the last century, this system changed from open, shifting sand dunes with sparse vegetation (including native beach grass, Elymus mollis), to densely vegetated continuous foredune ridges resulting from the introduction and subsequent invasions of two nonnative grass species (Ammophila arenaria and Ammophila breviligulata), each of which is associated with different dune shapes and sediment supply rates along the coast. Here we propose a biophysical feedback responsible for differences in dune shape, and we investigate two, non-mutually exclusive ecological mechanisms for these differences: (1) species differ in their ability to capture sand and (2) species differ in their growth habit in response to sand deposition. To investigate sand capture, we used a moveable bed wind tunnel experiment and found that increasing tiller density increased sand capture efficiency and that, under different experimental densities, the native grass had higher sand capture efficiency compared to the Ammophila congeners. However, the greater densities of nonnative grasses under field conditions suggest that they have greater potential to capture more sand overall. We used a mesocosm experiment to look at plant growth responses to sand deposition and found that, in response to increasing sand supply rates, A. arenaria produced higher-density vertical tillers (characteristic of higher sand capture efficiency), while A. breviligulata and E. mollis responded with lower-density lateral tiller growth (characteristic of lower sand capture efficiency). Combined, these experiments provide evidence for a species-specific effect on coastal dune shape. Understanding how dominant ecosystem engineers, especially nonnative ones, differ in their interactions with abiotic factors is necessary to better parameterize coastal vulnerability models and inform management practices related to both coastal protection ecosystem services and ecosystem restoration.     

Diversity-invasibility across an experimental disturbance gradient in Appalachian Forests

Research examining the relationship between community diversity and invasions by nonnative species has raised new questions about the theory and management of biological invasions. Ecological theory predicts, and small-scale experiments confirm, lower levels of nonnative species invasion into species-rich compared to species-poor communities, but observational studies across a wider range of scales often report positive relationships between native and nonnative species richness. This paradox has been attributed to the scale dependency of diversity-invasibility relationships and to differences between experimental and observational studies. Disturbance is widely recognized as an important factor determining invasibility of communities, but few studies have investigated the relative and interactive roles of diversity and disturbance on nonnative species invasion. Here, we report how the relationship between native and nonnative plant species richness responded to an experimentally applied disturbance gradient (from no disturbance up to clearcut) in oak-dominated forests. We consider whether results are consistent with various explanations of diversity-invasibility relationships including biotic resistance, resource availability, and the potential effects of scale (1 m2 to 2 ha). We found no correlation between native and nonnative species richness before disturbance except at the largest spatial scale, but a positive relationship after disturbance across scales and levels of disturbance. Post-disturbance richness of both native and nonnative species was positively correlated with disturbance intensity and with variability of residual basal area of trees. These results suggest that more nonnative plants may invade species-rich communities compared to species-poor communities following disturbance.     

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。     

Possibilities for management of coastal foredunes with deteriorated stands ofAmmophila arenaria (marram grass)

Ammophila arenaria (marram grass) is the most important plant species for sand stabilization in European coastal foredunes. Stand degeneration due to poor supply of wind-blown sand enhances the susceptibility for wind erosion when successional species do not become established. ReplantedA. arenaria often fails to become established. In the present study we examined whether management practices such as mowing, fertilizing, burning or below-ground cutting of plants may be applied to re-establish the vigour ofA. arenaria. Field experiments performed at exposed sites, where naturally succeeding plant species are not supposed to become established due to salt spray, showed that none of the applied methods resulted into enhanced tiller density ofA. arenaria. Thus, further studies are necessary to solve this type of management problem. At the leeward side of foredunes, the successional speciesFestuca rubra ssp.arenaria andElymus athericus could be established successfully as both pre-grown seedlings and planted bundles of cuttings (all further experiments were fertilized). When planted in spring, cuttings of successional plant species established less than those planted in early winter. Water repellency of the sand surface is supposed to be the main cause for this. Pre-grown seedlings were less susceptible for the season of planting. Direct sowing was not effective. Rabbit browsing had to be omitted to obtain successful establishment. When living rhizomes of successional plant species were still present in the soil profile, fencing alone turned out to be effective to re-establish vegetation at initially bare sites.     

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.     

Fish distributions and nutrient cycling in streams: can fish create biogeochemical hotspots?

Rates of biogeochemical processes often vary widely in space and time, and characterizing this variation is critical for understanding ecosystem functioning. In streams, spatial hotspots of nutrient transformations are generally attributed to physical and microbial processes. Here we examine the potential for heterogeneous distributions of fish to generate hotspots of nutrient recycling. We measured nitrogen (N) and phosphorus (P) excretion rates of 47 species of fish in an N-limited Neotropical stream, and we combined these data with population densities in each of 49 stream channel units to estimate unit- and reach-scale nutrient recycling. Species varied widely in rates of N and P excretion as well as excreted N:P ratios (6-176 molar). At the reach scale, fish excretion could meet >75% of ecosystem demand for dissolved inorganic N and turn over the ambient NH4 pool in <0.3 km. Areal N excretion estimates varied 47-fold among channel units, suggesting that fish distributions could influence local N availability. P excretion rates varied 14-fold among units but were low relative to ambient concentrations. Spatial variation in aggregate nutrient excretion by fish reflected the effects of habitat characteristics (depth, water velocity) on community structure (body size, density, species composition), and the preference of large-bodied species for deep runs was particularly important. We conclude that the spatial distribution of fish could indeed create hotspots of nutrient recycling during the dry season in this species-rich tropical stream. The prevalence of patchy distributions of stream fish and invertebrates suggests that hotspots of consumer nutrient recycling may often occur in stream ecosystems.