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Recent work has suggested that a tradeoff exists between habitat area and habitat heterogeneity, with a moderate amount of heterogeneity supporting greatest species richness. Support for this unimodal relationship has been mixed and has differed among habitats and taxa. We examined the relationship between habitat heterogeneity and species richness after accounting for habitat area in glacially formed wetlands in the Prairie Pothole Region in the United States at both local and landscape scales. We tested for area–habitat heterogeneity tradeoffs in wetland bird species richness, the richness of groups of similar species, and in species' abundances. We then identified the habitat relationships for individual species and the relative importance of wetland area vs. habitat heterogeneity and other wetland characteristics. We found that habitat area was the primary driver of species richness and abundance. Additional variation in richness and abundance could be explained by habitat heterogeneity or other wetland and landscape characteristics. Overall avian species richness responded unimodally to habitat heterogeneity, suggesting an area–heterogeneity tradeoff. Group richness and abundance metrics showed either unimodal or linear relationships with habitat heterogeneity. Habitat heterogeneity indices at local and landscape scales were important for some, but not all, species and avian groups. Both abundance of individual species and species richness of most avian groups were higher on publicly owned wetlands than on privately owned wetlands, on restored wetlands than natural wetlands, and on permanent wetlands than on wetlands of other classes. However, we found that all wetlands examined, regardless of ownership, restoration status, and wetland class, supported wetland-obligate birds. Thus, protection of all wetland types contributes to species conservation. Our results support conventional wisdom that protection of large wetlands is a priority but also indicate that maintaining habitat heterogeneity will enhance biodiversity and support higher populations of individual species.

Mature trees scattered throughout agricultural landscapes are critical habitat for some biota and provide a range of ecosystem services. These trees are declining in intensively managed agricultural landscapes globally. We developed a simulation model to predict the rates at which these trees are declining, identified the key variables that can be manipulated to mitigate this decline, and compared alternative management proposals. We used the initial numbers of trees in the stand, the predicted ages of these trees, their rate of growth, the number of recruits established, the frequency of recruitment, and the rate of tree mortality to simulate the dynamics of scattered trees in agricultural landscapes. We applied this simulation model to case studies from Spain, United States, Australia, and Costa Rica. We predicted that mature trees would be lost from these landscapes in 90-180 years under current management. Existing management recommendations for these landscapes--which focus on increasing recruitment--would not reverse this trend. The loss of scattered mature trees was most sensitive to tree mortality, stand age, number of recruits, and frequency of recruitment. We predicted that perpetuating mature trees in agricultural landscapes at or above existing densities requires a strategy that keeps mortality among established trees below around 0.5% per year, recruits new trees at a rate that is higher than the number of existing trees, and recruits new trees at a frequency in years equivalent to around 15% of the maximum life expectancy of trees. Numbers of mature trees in landscapes represented by the case studies will decline before they increase, even if strategies of this type are implemented immediately. This decline will be greater if a management response is delayed. /// Los árboles dispersos en paisajes agrícolas son hábitat critico para la biota y proporcionan una variedad de servicios ecológicos. Estos árboles están declinando globalmente en paisajes agrícolas manejados intensivamente. Desarrollamos un modelo de simulación para predecir las tasas a las que están declinando estos árboles, identificamos las principales variables que pueden ser manipuladas para mitigar esta declinación y comparamos propuestas de manejo alternativas. Utilizamos el número inicial de árboles en el sitio, las edades de estos árboles, su tasa de crecimiento, el número de individuos reclutados, la frecuencia de reclutamiento y la tasa de mortalidad de árboles para simular la dinámica de árboles dispersos en paisajes agrícolas. Aplicamos este modelo a estudios de caso de España, Estados Unidos, Australia y Costa Rica. Pronosticamos que los árboles maduros se perderán de estos paisajes entre 90 y 180 años bajos las condiciones de manejo actuales; las recomendaciones de manejo existentes - enfocadas en el incremento del reclutamiento - no cambiarían esta tendencia. Mediante la simulación de escenarios representando observaciones que abarcan todos los estudios de caso y una gama de opciones de manejo pudimos hacer recomendaciones genéricas sobre el manejo de árboles dispersos en paisajes agrícolas. La pérdida de árboles maduros dispersos fue más sensible a la mortalidad de árboles, edad del sitio, número de reclutas y frecuencia de reclutamiento. Predecimos que la perpetuación de árboles maduros en paisajes agrícolas en o por encima de las densidades existentes requiere de una estrategia que mantenga la mortalidad de árboles establecidos por debajo de 0.5% por año, que reclute árboles a una tasa mayor que el número de árboles existentes y reclute árboles nuevos en una frecuencia en años equivalente a alrededor de 15% de la esperanza de vida máxima de los árboles. Sin embargo, el número de árboles maduros en los paisajes representados por los estudios de caso declinará antes de incrementar, aun si estrategias de este tipo son implementadas inmediatamente. Esta declinación será mayor si se posterga una respuesta de manejo.

To anticipate the rapidly changing world resulting from global climate change, the projections of climate models must be incorporated into conservation. This requires that the scales of conservation be aligned with the scales of climate-change projections. We considered how conservation has incorporated spatial scale into protecting biodiversity, how the projections of climate-change models vary with scale, and how the two do or do not align. Conservation planners use information about past and current ecological conditions at multiple scales to identify conservation targets and threats and guide conservation actions. Projections of climate change are also made at multiple scales, from global and regional circulation models to projections downscaled to local scales. These downscaled projections carry with them the uncertainties associated with the broad-scale models from which they are derived; thus, their high resolution may be more apparent than real. Conservation at regional or global scales is about establishing priorities and influencing policy. At these scales, the coarseness and uncertainties of global and regional climate models may be less important than what they reveal about possible futures. At the ecoregional scale, the uncertainties associated with downscaling climate models become more critical because the distributions of conservation targets on which plans are founded may shift under future climates. At a local scale, variations in topography and land cover influence local climate, often overriding the projections of broad-scale climate models and increasing uncertainty. Despite the uncertainties, ecologists and conservationists must work with climate-change modelers to focus on the most likely projections. The future will be different from the past and full of surprises; judicious use of model projections at appropriate scales may help us prepare.

Despite striking differences in climate, soils, and evolutionary history among diverse biomes ranging from tropical and temperate forests to alpine tundra and desert, we found similar interspecific relationships among leaf structure and function and plant growth in all biomes. Our results thus demonstrate convergent evolution and global generality in plant functioning, despite the enormous diversity of plant species and biomes. For 280 plant species from two global data sets, we found that potential carbon gain (photosynthesis) and carbon loss (respiration) increase in similar proportion with decreasing leaf life-span, increasing leaf nitrogen concentration, and increasing leaf surface area-to-mass ratio. Productivity of individual plants and of leaves in vegetation canopies also changes in constant proportion to leaf life-span and surface area-to-mass ratio. These global plant functional relationships have significant implications for global scale modeling of vegetation-atmosphere CO2 exchange

For 76 annual, biennial, and perennial species common in the grasslands of central Minnesota, USA, we determined the patterns of correlations among seven organ-level traits (specific leaf area, leaf thickness, leaf tissue density, leaf angle, specific root length, average fine root diameter, and fine root tissue density) and their relationships with two traits relating to growth form (whether species existed for part of the growing season in basal, non-caulescent form and whether species were rhizomatous or not). The first correlation of traits showed that grasses had thin, dense leaves and thin roots while forbs had thick, low-density leaves and thick roots without any significant differences in growth form or life history. The second correlation of traits showed a gradient of species from those with high-density roots and high-density erect leaves to species with low-density roots and low-density leaves that were held parallel to the ground. High tissue density species were more likely to exist as a basal rosette for part of the season, were less likely to be rhizomatous, and less likely to be annuals. We examined the relationships between the two axes that represent the correlations of traits and previously collected data on the relative abundance of species across gradients of nitrogen addition and disturbance. Grasses were generally more abundant than forbs and the relative abundance of grasses and forbs did not change with increasing nitrogen addition or soil disturbance. High tissue density species became less common as fertility and disturbance increased.

Species diversity has declined in ecosystems worldwide as a result of habitat fragmentation, eutrophication, and land-use change. If such decline is to be halted ecological mechanisms that restore or maintain biodiversity are needed. Two long-term field experiments were performed in native grassland to assess the effects of fire, nitrogen addition, and grazing or mowing on plant species diversity. In one experiment, richness declined on burned and fertilized treatments, whereas mowing maintained diversity under these conditions. In the second experiment, loss of species diversity due to frequent burning was reversed by bison, a keystone herbivore in North American grasslands. Thus, mowing or the reestablishment of grazing in anthropogenically stressed grasslands enhanced biodiversity

Prior to European colonization and the introduction of cattle, bison were the prominent grazing ungulates throughout North American grasslands. Yet, relative zoogeomorphic impacts between bison and cattle on grassland streams remain largely unknown. Utilizing a paired watershed study design, we compared baseflow suspended sediment concentrations across ten watersheds and four grazing treatments (ungrazed, bison-grazed, moderately stocked cattle-grazed, and intensively stocked cattle-grazed) in the Flint Hills subregion of the Great Plains. Additionally, we determined whether periods of increased thermal stress led to higher sediment concentrations within each treatment. Water samples were analyzed for total suspended solids (TSS, mg/L), nonvolatile suspended solids (NVSS, mg/L), and percentage organic matter (POM, percent). Intensively stocked cattle-grazed treatments produced significantly higher TSS and NVSS concentrations relative to ungrazed (TSS p = 0.012, NVSS p < 0.01) and bison-grazed treatments (TSS p = 0.082, NVSS p < 0.01). Moderately stocked cattle-grazed treatments contained significantly higher NVSS concentrations relative to bison-grazed treatments (p = 0.057). Bison-grazed and ungrazed treatments contained similar sediment concentrations (TSS and NVSS p > 0.10). Additionally, intensively and moderately stocked cattle-grazed treatments showed a significant increase in sediment concentrations with increasing temperature (p = 0.024 and p = 0.08, respectively), whereas bison-grazed and ungrazed treatments did not (p > 0.10). At the subdaily timescale, the highest sediment concentrations within cattle-grazed treatments and the greatest difference in sediment concentrations between cattle-grazed and ungrazed treatments coincided with the hottest daily temperatures, further highlighting that cattle-grazing impacts are influenced by thermal conditions.

This paper examines four key mechanisms of the seed size/number trade-off (SSNT) models to assess their relevance to a general understanding of plant community structure. Mechanism 1 is that large seeds have a greater probability of winning in competition against smaller seeds. I provide interspecific experimental evidence that there is a competitive hierarchy among seedlings based on seed size. Mechanism 2 is that a trade-off exists between the number and size of seeds produced for a given reproductive allocation. Negative correlations between seed size and number were found consistently across a range of species from a range of habitats, from published literature. Mechanism 3, that seedling-seedling competition is an important influence on species composition, was found to exist potentially in a range of environments, including annual-dominated, post-fire and gap-dynamic communities. However, there is little quantitative evidence available and this is likely to be a restrictive mechanism. Mechanism 4, that small seeds are superior colonists due to their greater number, was tested in a field experiment in a calcareous grassland community. No supporting evidence was found, suggesting that the SSNT is not an important determinant of structure in this community. Thus two of the four mechanisms can be considered to hold true generally, while the third mechanism may be valid in particular environments. The fourth mechanism did not apply in the community tested, but could be tested in a wider range of communities.

Some theories and experimental studies suggest that areas of low plant species richness may be invaded more easily than areas of high plant species richness. We gathered nested-scale vegetation data on plant species richness, foliar cover, and frequency from 200 1-m2subplots (20 1000-m2modified-Whittaker plots) in the Colorado Rockies (USA), and 160 1-m2subplots (16 1000-m2plots) in the Central Grasslands in Colorado, Wyoming, South Dakota, and Minnesota (USA) to test the generality of this paradigm. At the 1-m2scale, the paradigm was supported in four prairie types in the Central Grasslands, where exotic species richness declined with increasing plant species richness and cover. At the 1-m2scale, five forest and meadow vegetation types in the Colorado Rockies contradicted the paradigm; exotic species richness increased with native-plant species richness and foliar cover. At the 1000-m2plot scale (among vegetation types), 83% of the variance in exotic species richness in the Central Grasslands was explained by the total percentage of nitrogen in the soil and the cover of native plant species. In the Colorado Rockies, 69% of the variance in exotic species richness in 1000-m2plots was explained by the number of native plant species and the total percentage of soil carbon. At landscape and biome scales, exotic species primarily invaded areas of high species richness in the four Central Grasslands sites and in the five Colorado Rockies vegetation types. For the nine vegetation types in both biomes, exotic species cover was positively correlated with mean foliar cover, mean soil percentage N, and the total number of exotic species. These patterns of invasibility depend on spatial scale, biome and vegetation type, spatial autocorrelation effects, availability of resources, and species-specific responses to grazing and other disturbances. We conclude that: (1) sites high in herbaceous foliar cover and soil fertility, and hot spots of plant diversity (and biodiversity), are invasible in many landscapes; and (2) this pattern may be more closely related to the degree resources are available in native plant communities, independent of species richness. Exotic plant invasions in rare habitats and distinctive plant communities pose a significant challenge to land managers and conservation biologists.

Almost all natural plant communities contain arbuscular mycorrhizal fungi (AMF). We hypothesized that the species composition of AMF communities could have the potential to determine plant community structure if the growth response to different AMF species or to communities of AMF species varies among plant species. To test the existence of such a differential response we conducted a pot experiment where each of three plant species, Hieracium pilosella, Bromus erectus, and Festuca ovina were inoculated with each of four AMF species, or with a mixture of these four AMF species, or were uninoculated. The AMF species originated from a calcareous grassland in which the three plant species also coexisted. We obtained three pieces of evidence suggesting that AMF have the potential to determine plant community structure. First, plant species differed in their dependency on AMF, thus varying in degree of benefit received. Second, specific AMF species and a mixture of these AMF species had significantly different effects on several plant growth variables, and these effects were not the same on each plant species. Third, the amount of variation in the growth response of a plant species to four AMF species and to the mixture of AMF species differed among the plant species. Hieracium differed greatly in its growth response to several AMF species while Bromus did not exhibit much variation in its response to different AMF species. The varying mycorrhizal dependency of different plant species has previously been proposed as a mechanism determining plant community structure. However, we found that the mycorrhizal dependency of a plant species can vary greatly because of differential growth responses to specific AMF species compared to the growth of the uninoculated plants. Consequently mycorrhizal dependency, as a measure indicating how much a plant depends on AMF for its growth, is not necessarily a fixed value and therefore cannot be used as a definitive term. In addition, those plant species with highly variable responses to single AMF species or to combinations of AMF species (AMF communities) will be strongly affected by the specific species of AMF that occupy their roots, in contrast to plant species that do not respond differently to different AMF species. We conclude that, through their differential effects on plant growth, AMF species that co-occur as natural AMF communities have the potential to determine plant community structure, and that future studies on plant population and community structure need to consider the strength of their role as a determinant.