Feces nitrogen release induced by different large herbivores in a dry grassland.

Large herbivores have pronounced effects on nutrient cycling in grasslands. These organisms are known to alter the quality and quantity of plant production as well as the amounts and quality of plant litter and animal wastes. The generalization that the relative quality of detritus inputs is enhanced by herbivores is well known, but how this process is affected by diet selection processing and feces production of different large herbivores remains largely unstudied. Here, we measured how these differences for cattle and sheep on a dry grassland might influence nitrogen (N) mineralization from feces. We found that cattle of larger body size tended to select the low quality grass Stipa grandis as their major food source. In contrast, the subdominant grass Leymus chinensis, with relatively high N content, was a majority in the diet of smaller sheep, when palatable forbs were insufficient in the field. This diverse diet quality resulted in a C:N ratio of cattle feces that was higher than that of sheep feces. Relatively higher labile C availability in the cattle feces, namely relatively higher cellulose/hemicellulose contents, promoted microbial growth and in turn accelerated cattle feces decomposition. A surprise finding was that the feces from cattle mineralized about twice as much N as feces from sheep, despite the latter having slightly higher N content. From a grassland productivity perspective, increasing the proportion of large body-sized species in grazing herbivore assemblages perhaps is beneficial to forage productivity and nutrient recycling by the rapid degradation of feces.

Priorities for research in soil ecology.

The ecological interactions that occur in and with soil are of consequence in many ecosystems on the planet. These interactions provide numerous essential ecosystem services, and the sustainable management of soils has attracted increasing scientific and public attention. Although soil ecology emerged as an independent field of research many decades ago, and we have gained important insights into the functioning of soils, there still are fundamental aspects that need to be better understood to ensure that the ecosystem services that soils provide are not lost and that soils can be used in a sustainable way. In this perspectives paper, we highlight some of the major knowledge gaps that should be prioritized in soil ecological research. These research priorities were compiled based on an online survey of 32 editors of Pedobiologia - Journal of Soil Ecology. These editors work at universities and research centers in Europe, North America, Asia, and Australia.The questions were categorized into four themes: (1) soil biodiversity and biogeography, (2) interactions and the functioning of ecosystems, (3) global change and soil management, and (4) new directions. The respondents identified priorities that may be achievable in the near future, as well as several that are currently achievable but remain open. While some of the identified barriers to progress were technological in nature, many respondents cited a need for substantial leadership and goodwill among members of the soil ecology research community, including the need for multi-institutional partnerships, and had substantial concerns regarding the loss of taxonomic expertise.

Plant community and soil chemistry responses to long-term nitrogen inputs drive changes in alpine bacterial communities.

Bacterial community composition and diversity was studied in alpine tundra soils across a plant species and moisture gradient in 20 y-old experimental plots with four nutrient addition regimes (control, nitrogen (N), phosphorus (P) or both nutrients). Different bacterial communities inhabited different alpine meadows, reflecting differences in moisture, nutrients and plant species. Bacterial community alpha-diversity metrics were strongly correlated with plant richness and the production of forbs. After meadow type, N addition proved the strongest determinant of bacterial community structure. Structural Equation Modeling demonstrated that tundra bacterial community responses to N addition occur via changes in plant community composition and soil pH resulting from N inputs, thus disentangling the influence of direct (resource availability) vs. indirect (changes in plant community structure and soil pH) N effects that have remained unexplored in past work examining bacterial responses to long-term N inputs in these vulnerable environments. Across meadow types, the relative influence of these indirect N effects on bacterial community structure varied. In explicitly evaluating the relative importance of direct and indirect effects of long-term N addition on bacterial communities, this study provides new mechanistic understandings of the interaction between plant and microbial community responses to N inputs amidst environmental change.

Imposing antecedent global change conditions rapidly alters plant community composition in a mixed-grass prairie.

Global change drivers are altering climatic and edaphic conditions of ecosystems across the globe, and we expect novel plant communities to become more common as a result. In the Colorado Front Range, compositional changes have occurred in the mixed-grass prairie plant community in conjunction with shifts in winter precipitation and atmospheric nitrogen (N) deposition. To test whether these environmental changes have been responsible for the observed plant community change, we conducted an in situ manipulative experiment in a mixed-grass meadow near Boulder, CO. We simulated historical conditions by reducing N availability (+500 g C m(-2) year(-1)) and winter precipitation (with rainout shelters) for 2 years (2013-2014) and compared vegetation response to these treatments with that of ambient conditions. The site experienced an extreme precipitation event in autumn 2013 which allowed comparison of an exceptionally wet year with an average year. We measured pre-treatment species composition in 2012, and treatment responses in the spring and summer of 2013 and 2014. As predicted, simulating historical low N-winter dry conditions resulted in a plant community dominated by historically abundant species. Cool-season introduced species were significantly reduced in low N-winter dry plots, particularly the annual plants Bromus tectorum and Alyssum parviflorum. These same species responded strongly to the extreme precipitation event with large increases, while native grasses and forbs showed little change in productivity or composition under varying climatic or edaphic conditions. This work provides clear evidence linking on-going global change drivers to altered plant community composition in an otherwise relatively undisturbed grassland ecosystem.

Effects of precipitation change and neighboring plants on population dynamics of Bromus tectorum.

Shifting precipitation patterns resulting from global climate change will influence the success of invasive plant species. In the Front Range of Colorado, Bromus tectorum (cheatgrass) and other non-native winter annuals have invaded grassland communities and are becoming more abundant. As the global climate warms, more precipitation may fall as rain rather than snow in winter, and an increase in winter rain could benefit early-growing winter annuals, such as B. tectorum, to the detriment of native species. In this study we measured the effects of simulated changes in seasonal precipitation and presence of other plant species on population growth of B. tectorum in a grassland ecosystem near Boulder, Colorado, USA. We also performed elasticity analyses to identify life transitions that were most sensitive to precipitation differences. In both study years, population growth rates were highest for B. tectorum growing in treatments receiving supplemental winter precipitation and lowest for those receiving the summer drought treatment. Survival of seedlings to flowering and seed production contributed most to population growth in all treatments. Biomass of neighboring native plants was positively correlated with reduced population growth rates of B. tectorum. However, exotic plant biomass had no effect on population growth rates. This study demonstrates how interacting effects of climate change and presence of native plants can influence the population growth of an invasive species. Overall, our results suggest that B. tectorum will become more invasive in grasslands if the seasonality of precipitation shifts towards wetter winters and allows B. tectorum to grow when competition from native species is low.

Biological control of invasive plant species: a reassessment for the Anthropocene.

The science of finding, testing and releasing herbivores and pathogens to control invasive plant species has achieved a level of maturity and success that argues for continued and expanded use of this program. The practice, however, remains unpopular with some conservationists, invasion biologists, and stakeholders. The ecological and economic benefits of controlling densities of problematic plant species using biological control agents can be quantified, but the risks and net benefits of biological control programs are often derived from social or cultural rather than scientific criteria. Management of invasive plants is a 'wicked problem', and local outcomes to wicked problems have both positive and negative consequences differentially affecting various groups of stakeholders. The program has inherent uncertainties; inserting species into communities that are experiencing directional or even transformational changes can produce multiple outcomes due to context-specific factors that are further confounded by environmental change drivers. Despite these uncertainties, biological control could play a larger role in mitigation and adaptation strategies used to maintain biological diversity as well as contribute to human well-being by protecting food and fiber resources.

The effects of black-tailed prairie dogs on plant communities within a complex urban landscape: an ecological surprise?

Historically, prairie dogs (Cynomys spp.) have been considered essential keystone species of western United States grassland ecosystems because they provide unique services and increase vegetation community richness, evenness, and diversity. However, the effects of black-tailed prairie dogs (Cynomys ludovicianus) on lands adjacent to or surrounded by urban areas may not result in the same ecosystem benefits historically associated with their presence. An urban landscape presents prairie dogs with movement challenges unparalleled in natural landscapes, as well as suites of nonnative plant species that are more common in disturbed areas. This study examined a complex ecosystem where vegetation communities are being influenced by directional environmental change, and quantified the synergistic effects resulting from the protective management of a native keystone species. The data set for this analysis was comprised of 71 paired (occupied by prairie dogs vs. unoccupied) vegetation surveys and 156 additional unpaired surveys collected from around the city of Boulder, Colorado, USA for 14 yr. Linear mixed models were used to compare data from transects occupied and unoccupied by prairie dogs, as well as to evaluate the effect of prairie dog occupation duration. In the absence of prairie dogs, vegetation in this region exhibited declines in native grasses, no changes in introduced grasses, and increases in native and nonnative forbs and bare soil over the study interval. In the presence of prairie dogs, these observed directional changes were nearly all amplified at rates four to 10 times greater than when prairie dogs were absent. Areas in Boulder occupied by prairie dogs also had significantly lower richness, evenness, and diversity of plant species, compared to unoccupied areas. Analysis of plant functional groups revealed the significant reduction of perennial native grasses, as well as a significantly higher cover of introduced forbs in occupied areas. Prairie dogs amplified the effects of low-impact environmental directional changes, creating more novel vegetation communities than the environmental factors alone. Results from this research provide insight regarding the synergistic effects of a native keystone species within a complex human-impacted 21st century landscape.

Nitrogen enrichment differentially affects above- and belowground plant defense.

PREMISE OF THE STUDY: Human nitrogen (N) inputs to terrestrial ecosystems have greatly increased in recent years and may have important consequences for plant growth, reproduction, and defense. Although numerous studies have investigated the effects of nitrogen addition on plants, few have examined both above- and belowground responses within a range of predicted increase and apart from concomitant increases in other nutrients. • METHODS: We conducted a greenhouse experiment to study the consequences of increased nitrogen inputs, such as those from atmospheric N deposition, on plant performance, chemical defenses, and allocation tradeoffs for an invasive species, Linaria dalmatica. This plant produces iridoid glycosides, which are a group of terpenoid compounds. • KEY RESULTS: Soil nitrogen enrichment increased growth, reproduction, and whole-plant iridoid glycosides while decreasing some costs of defense. Interestingly, nitrogen addition had varying effects on defense allocation to above- and belowground tissues. Specifically, there was no change in iridoid glycoside concentrations of shoots, whereas concentrations decreased in flowers by ~35% and increased in roots by >400%. • CONCLUSIONS: Observed increases in plant performance and chemical defenses may have implications for the invasion potential of L. dalmatica. Moreover, our results highlight the importance of evaluating both above- and belowground plant defenses. In particular, findings presented here indicate that research focused on leaf-level defenses may not detect key allelochemical responses, including changes in plant resistance traits that could affect consumers (e.g., herbivores and pathogens) that specialize on different plant tissues as well as plant fitness and invasion success.

Reconciling contradictory findings of herbivore impacts on spotted knapweed (Centaurea stoebe) growth and reproduction.

Substantial controversy surrounds the efficacy of biological control insects to reduce densities of Centaurea stoebe, a widespread, aggressive invasive plant in North America. We developed a graphical model to conceptualize the conditions required to explain the current contradictory findings, and then employed a series of manipulations to evaluate C. stoebe responses to herbivores. We manipulated soil nitrogen and competition in a field population and measured attack rates of a foliage and seed feeder (Larinus minutus), two gall flies (Urophora spp.), and a root feeder (Cyphocleonus achates), as well as their effects on the growth and reproduction of C. stoebe. Nitrogen limitation and competing vegetation greatly reduced C. stoebe growth. L. minutus most intensively reduced seed production in low-nitrogen soils, and removal of neighboring vegetation increased Larinus numbers per flower head and the percentage of flowers attacked by 15% and 11%, respectively. Cyphocleonus reduced flower production and aboveground biomass over two years, regardless of resources or competition. Our results, in conjunction with other published studies, demonstrate that positive, neutral, and negative plant growth responses to herbivory can be generated. However, under realistic field conditions and in the presence of multiple herbivores, our work repudiates earlier studies that indicate insect herbivores increase C. stoebe dominance.

Additive effects of aboveground and belowground herbivores on the dominance of spotted knapweed (Centaurea stoebe).

Spotted knapweed (Centaurea stoebe) is found in over 3 million ha of rangeland and forests across North America, and evidence supporting the use of biological control as a regional method to reduce infestations and their associated impacts remains inconclusive. Several species of insects have been reported to reduce plant densities in some areas; however, rigorous studies that test combinations of these species and the influence of resource availability are lacking. We examined the singular and combined effects of herbivory by a root weevil (Cyphocleonus achates) and a flower head weevil (Larinus minutus) on the growth and flower production of C. stoebe. We also manipulated soil resource fertility as an additional factor that could explain the outcomes of contradictory biological control herbivore effects on C. stoebe. In a greenhouse study, herbivory by C. achates decreased flower production for plants across all resource environments. In a caged common garden study, the negative effects of herbivory also did not interact with soil nutrient status. However, the presence of plant competition further decreased knapweed growth, and the negative effects of concurrent herbivory by C. achates and L. minutus on plant biomass and flower production were additive. Derived within the context of variable levels of soil nutrient availability and competing vegetation, these results support the cumulative stress hypothesis and the contention that combined above- and belowground herbivory can reduce spotted knapweed densities and reduce the ecological and economic impacts of this species in rangelands of western North America.

Competitive impacts and responses of an invasive weed: dependencies on nitrogen and phosphorus availability.

Changes in competitive interactions under conditions of enhanced resource availability could explain the invasion success of some problematic plant species. For one invader of North American grasslands, Centaurea diffusa (diffuse knapweed), we test three hypotheses: (1) under ambient (high resource) conditions, C. diffusa is better able to tolerate competition from the resident community (competitive response), (2) under ambient conditions, C. diffusa strong impacts the competitive environment (competitive effect), and (3) reductions in nitrogen and/or phosphorus availability diminish these advantages. In support of our first hypothesis, C. diffusa was the most tolerant to neighbor competition of the four focal species under current resource conditions. In opposition to our second hypothesis, however, neighborhoods that contained C. diffusa and those where C. diffusa had been selectively removed did not differ in their impact on the performance of target transplant individuals or on resource conditions. Reduction in resource availability influenced competitive tolerance but not competitive impact, in partial support of our last hypothesis. Reduction in soil nitrogen (via sucrose carbon addition) enhanced the degree of neighbor competition experienced by all species but did not change their relative rankings; C. diffusa remained the best competitor under low nitrogen conditions. Reduction of soil phosphorus (via gypsum addition) weakened the ability of C. diffusa to tolerate neighbor competition proportionately more than the other focal species. Consequently, under low phosphorus conditions, C. diffusa lost its competitive advantage and tolerated neighbor competition similarly to the other focal species. We conclude that C. diffusa invasion may be double-edged: C. diffusa is less limited by nitrogen than the other focal species and is better able to utilize phosphorus to its competitive advantage.

Soil ecological interactions: comparisons between tropical and subalpine forests.

Soil fauna can influence soil processes through interactions with the microbial community. Due to the complexity of the functional roles of fauna and their effects on microbes, little consensus has been reached on the extent to which soil fauna can regulate microbial activities. We quantified soil microbial biomass and maximum growth rates in control and fauna-excluded treatments in dry and wet tropical forests and north- and south-facing subalpine forests to test whether soil fauna effects on microbes are different in tropical and subalpine forests. Exclusion of fauna was established by physically removing the soil macrofauna and applying naphthalene. The effect of naphthalene application on the biomass of microbes that mineralize salicylate was quantified using the substrate induced growth response method. We found that: (1) the exclusion of soil fauna resulted in a higher total microbial biomass and lower maximum growth rate in the subalpine forests, (2) soil fauna exclusion did not affect the microbial biomass and growth rate in the tropical forests, and (3) the microbial biomass of salicylate mineralizers was significantly enhanced in the fauna-exclusion treatment in the tropical wet and the south-facing subalpine forests. We conclude that non-target effects of naphthalene on the microbial community alone cannot explain the large differences in total microbial biomass found between control and fauna-excluded treatments in the subalpine forests. Soil fauna have relatively larger effects on the microbial activities in the subalpine forests than in tropical dry and wet forests.