Showing papers in "Ecology in 2010"

A distance-based framework for measuring functional diversity from multiple traits

Abstract: A new framework for measuring functional diversity (FD) from multiple traits has recently been proposed. This framework was mostly limited to quantitative traits without missing values and to situations in which there are more species than traits, although the authors had suggested a way to extend their framework to other trait types. The main purpose of this note is to further develop this suggestion. We describe a highly flexible distance-based framework to measure different facets of FD in multidimensional trait space from any distance or dissimilarity measure, any number of traits, and from different trait types (i.e., quantitative, semi-quantitative, and qualitative). This new approach allows for missing trait values and the weighting of individual traits. We also present a new multidimensional FD index, called functional dispersion (FDis), which is closely related to Rao's quadratic entropy. FDis is the multivariate analogue of the weighted mean absolute deviation (MAD), in which the weights are species relative abundances. For unweighted presence-absence data, FDis can be used for a formal statistical test of differences in FD. We provide the "FD" R language package to easily implement our distance-based FD framework.

Disturbance and landscape dynamics in a changing world

Abstract: Disturbance regimes are changing rapidly, and the consequences of such changes for ecosystems and linked social-ecological systems will be profound. This paper synthesizes current understanding of disturbance with an emphasis on fundamental contributions to contemporary landscape and ecosystem ecology, then identifies future research priorities. Studies of disturbance led to insights about heterogeneity, scale, and thresholds in space and time and catalyzed new paradigms in ecology. Because they create vegetation patterns, disturbances also establish spatial patterns of many ecosystem processes on the landscape. Drivers of global change will produce new spatial patterns, altered disturbance regimes, novel trajectories of change, and surprises. Future disturbances will continue to provide valuable opportunities for studying pattern-process interactions. Changing disturbance regimes will produce acute changes in ecosystems and ecosystem services over the short (years to decades) and long-term (centuries and beyond). Future research should address questions related to (1) disturbances as catalysts of rapid ecological change, (2) interactions among disturbances, (3) relationships between disturbance and society, especially the intersection of land use and disturbance, and (4) feedbacks from disturbance to other global drivers. Ecologists should make a renewed and concerted effort to understand and anticipate the causes and consequences of changing disturbance regimes.

Functional traits and the growth–mortality trade-off in tropical trees

Abstract: A trade-off between growth and mortality rates characterizes tree species in closed canopy forests. This trade-off is maintained by inherent differences among species and spatial variation in light availability caused by canopy-opening disturbances. We evaluated conditions under which the trade-off is expressed and relationships with four key functional traits for 103 tree species from Barro Colorado Island, Panama. The trade-off is strongest for saplings for growth rates of the fastest growing individuals and mortality rates of the slowest growing individuals (r 2 ¼ 0.69), intermediate for saplings for average growth rates and overall mortality rates (r 2 ¼ 0.46), and much weaker for large trees (r 2 � 0.10). This parallels likely levels of spatial variation in light availability, which is greatest for fast- vs. slow-growing saplings and least for large trees with foliage in the forest canopy. Inherent attributes of species contributing to the trade-off include abilities to disperse, acquire resources, grow rapidly, and tolerate shade and other stresses. There is growing interest in the possibility that functional traits might provide insight into such ecological differences and a growing consensus that seed mass (SM), leaf mass per area (LMA), wood density (WD), and maximum height (Hmax) are key traits among forest trees. Seed mass, LMA, WD, and Hmax are predicted to be small for light-demanding species with rapid growth and mortality and large for shade-tolerant species with slow growth and mortality. Six of these trait-demographic rate predictions were realized for saplings; however, with the exception of WD, the relationships were weak (r 2 , 0.1 for three and r 2 , 0.2 for five of the six remaining relationships). The four traits together explained 43-44% of interspecific variation in species positions on the growth-mortality trade-off; however, WD alone accounted for .80% of the explained variation and, after WD was included, LMA and Hmax made insignificant contributions. Virtually the full range of values of SM, LMA, and Hmax occurred at all positions on the growth-mortality trade-off. Although WD provides a promising start, a successful trait- based ecology of tropical forest trees will require consideration of additional traits.

Abandoning Sverdrup's Critical Depth Hypothesis on phytoplankton blooms

Abstract: The Critical Depth Hypothesis formalized by Sverdrup in 1953 posits that vernal phytoplankton blooms occur when surface mixing shoals to a depth shallower than a critical depth horizon defining the point where phytoplankton growth exceeds losses. This hypothesis has since served as a cornerstone in plankton ecology and reflects the very common assumption that blooms are caused by enhanced growth rates in response to improved light, temperature, and stratification conditions, not simply correlated with them. Here, a nine-year satellite record of phytoplankton biomass in the subarctic Atlantic is used to reevaluate seasonal plankton dynamics. Results show that (1) bloom initiation occurs in the winter when mixed layer depths are maximum, not in the spring, (2) coupling between phytoplankton growth (l) and losses increases during spring stratification, rather than decreases, (3) maxima in net population growth rates (r) are as likely to occur in midwinter as in spring, and (4) r is generally inversely related to l. These results are incompatible with the Critical Depth Hypothesis as a functional framework for understanding bloom dynamics. In its place, a ''Dilution-Recoupling Hypothesis'' is described that focuses on the balance between phytoplankton growth and grazing, and the seasonally varying physical processes influencing this balance. This revised view derives from fundamental concepts applied during field dilution experiments, builds upon earlier modeling results, and is compatible with observed phytoplankton blooms in the absence of spring mixed layer shoaling.

The ecology of climate change and infectious diseases: reply

Abstract: Paul Epstein (2010) objects to my hypothesis that climate change should result in shifts (as opposed to net increases) in the distribution of infectious diseases (Lafferty 2009a, b). My first exposure to the link between climate change and infectious disease came from a Scientific American article by Epstein (2000) showing maps of malaria transmission risk increasing dramatically in New England and Europe. I used these maps to frighten my North American students into caring about malaria. In 2005, I attended a Cary Conference where David Rogers and Sarah Randolph challenged Epstein’s view that climate change would unilaterally increase disease. They advocated two-tailed hypotheses (disease can go up or down), followed by sophisticated analyses that control for temporal and spatial confounds and apportion variance in disease to climate and myriad other factors. Those talks convinced me ecologists possessed the skills needed to address this important issue and that a review aimed at ecologists might help get them into the game. Epstein gives two arguments for why disease should respond positively (but not negatively) to climate change. The first considers that the future might have warmer winters and disproportionate warming at higher latitudes. As a result, the globe would not necessarily be hotter than at present, just less cold. The tropics would stay the same and the temperate latitudes would become

Functional traits and environmental filtering drive community assembly in a species-rich tropical system

Abstract: Mechanistic models of community assembly state that biotic and abiotic filters constrain species establishment through selection on their functional traits. Predicting this assembly process is hampered because few studies directly incorporate environmental measurements and scale up from species to community level and because the functional traits' significance is environment dependent. We analyzed community assembly by measuring structure, environmental conditions, and species traits of secondary forests in a species-rich tropical system. We found, as hypothesized, that community structure shaped the local environment and that strong relationships existed between this environment and the traits of the most successful species of the regeneration communities. Path and multivariate analyses showed that temperature and leaf traits that regulate it were the most important factors of community differentiation. Comparisons between the trait composition of the forest's regeneration, juvenile, and adult communities showed a consistent community assembly pattern. These results allowed us to identify the major functional traits and environmental factors involved in the assembly of dry-forest communities and demonstrate that environmental filtering is a predictable and fundamental process of community assembly, even in a complex system such as a tropical forest.

Consistent effects of nitrogen fertilization on soil bacterial communities in contrasting systems

Abstract: Ecosystems worldwide are receiving increasing amounts of reactive nitrogen (N) through anthropogenic activities. Although the effects of increased N inputs on plant communities have been reasonably well studied, few comparable studies have examined impacts on whole soil bacterial communities, though they play critical roles in ecosystem functioning. We sampled soils from two long-term ecological research (LTER) experimental N gradients, both of which have been amended with NH4NO3; a grassland at Cedar Creek (27 years of N additions) and an agricultural field at Kellogg Biological Station (8 years of N additions). By examining shifts in bacterial communities across these contrasting ecosystem types, we could test competing hypotheses about the direct and indirect factors that might drive bacterial responses to elevated N inputs. Bacterial community structure was highly responsive to N additions. We observed predictable and consistent changes in the structure of the bacterial communities across both ecosystem types. Our results suggest that bacterial communities across these gradients are more structured by N and/or soil carbon availability than by shifts in the plant community or soil pH associated with the elevated nitrogen inputs. In contrast to the pronounced shifts in bacterial community composition and in direct contrast to the patterns often observed in plant communities, increases in N availability did not have consistent effects on the richness and diversity of soil bacterial communities.

General stabilizing effects of plant diversity on grassland productivity through population asynchrony and overyielding

Abstract: Insurance effects of biodiversity can stabilize the functioning of multispecies ecosystems against environmental variability when differential species' responses lead to asynchronous population dynamics. When responses are not perfectly positively correlated, declines in some populations are compensated by increases in others, smoothing variability in ecosystem productivity. This variance reduction effect of biodiversity is analogous to the risk-spreading benefits of diverse investment portfolios in financial markets. We use data from the BIODEPTH network of grassland biodiversity experiments to perform a general test for stabilizing effects of plant diversity on the temporal variability of individual species, functional groups, and aggregate communities. We tested three potential mechanisms: reduction of temporal variability through population asynchrony; enhancement of long-term average performance through positive selection effects; and increases in the temporal mean due to overyielding. Our results support a stabilizing effect of diversity on the temporal variability of grassland aboveground annual net primary production through two mechanisms. Two-species communities with greater population asynchrony were more stable in their average production over time due to compensatory fluctuations. Overyielding also stabilized productivity by increasing levels of average biomass production relative to temporal variability. However, there was no evidence for a performance-enhancing effect on the temporal mean through positive selection effects. In combination with previous work, our results suggest that stabilizing effects of diversity on community productivity through population asynchrony and overyielding appear to be general in grassland ecosystems.

Plant diversity effects on soil microorganisms support the singular hypothesis.

Abstract: The global decline in biodiversity has generated concern over the consequences for ecosystem functioning and services. Although ecosystem functions driven by soil microorganisms such as plant productivity, decomposition, and nutrient cycling are of particular importance, interrelationships between plant diversity and soil microorganisms are poorly understood. We analyzed the response of soil microorganisms to variations in plant species richness (1-60) and plant functional group richness (1-4) in an experimental grassland system over a period of six years. Major abiotic and biotic factors were considered for exploring the mechanisms responsible for diversity effects. Further, microbial growth characteristics were assessed following the addition of macronutrients. Effects of plant diversity on soil microorganisms were most pronounced in the most diverse plant communities though differences only became established after a time lag of four years. Differences in microbial growth characteristics indicate successional changes from a disturbed (zymogeneous) to an established (autochthonous) microbial community four years after establishment of the experiment. Supporting the singular hypothesis for plant diversity, the results suggest that plant species are unique, each contributing to the functioning of the belowground system. The results reinforce the need for long-term biodiversity experiments to fully appreciate consequences of current biodiversity loss for ecosystem functioning.

Priming effect: bridging the gap between terrestrial and aquatic ecology

Abstract: Understanding how ecosystems store or release carbon is one of ecology's greatest challenges in the 21st century. Organic matter covers a large range of chemical structures and qualities, and it is classically represented by pools of different recalcitrance to degradation. The interaction effects of these pools on carbon cycling are still poorly understood and are most often ignored in global-change models. Soil scientists have shown that inputs of labile organic matter frequently tend to increase, and often double, the mineralization of the more recalcitrant organic matter. The recent revival of interest for this phenomenon, named the priming effect, did not cross the frontiers of the disciplines. In particular, the priming effect phenomenon has been almost totally ignored by the scientific communities studying marine and continental aquatic ecosystems. Here we gather several arguments, experimental results, and field observations that strongly support the hypothesis that the priming effect is a general phenomenon that occurs in various terrestrial, freshwater, and marine ecosystems. For example, the increase in recalcitrant organic matter mineralization rate in the presence of labile organic matter ranged from 10% to 500% in six studies on organic matter degradation in aquatid ecosystems. Consequently, the recalcitrant organic matter mineralization rate may largely depend on labile organic matter availability, influencing the CO2 emissions of both aquatic and terrestrial ecosystems. We suggest that (1) recalcitrant organic matter may largely contribute to the CO2 emissions of aquatic ecosystems through the priming effect, and (2) priming effect intensity may be modified by global changes, interacting with eutrophication processes and atmospheric CO2 increases. Finally, we argue that the priming effect acts substantially in the carbon and nutrient cycles in all ecosystems. We outline exciting avenues for research, which could provide new insights on the responses of ecosystems to anthropogenic perturbations and their feedbacks to climatic changes.

Quantifying Bufo boreas connectivity in Yellowstone National Park with landscape genetics

Abstract: A major objective of ecology is to understand how ecological processes limit population connectivity and species' distributions. By spatially quantifying ecological components driving functional connectivity, we can understand why some locally suitable habitats are unoccupied, resulting in observed discontinuities in distribution. However, estimating connectivity may be difficult due to population stochasticity and violations of assumptions of parametric statistics. To address these issues, we present a novel application of Random Forests to landscape genetic data. We address the effects of three key ecological components on Bufo boreas connectivity in Yellowstone National Park: ecological process, scale, and hierarchical organization. Habitat permeability, topographic morphology, and temperature-moisture regime are all significant ecological processes associated with B. boreas connectivity. Connectivity was influenced by growing-season precipitation, 1988 Yellowstone fires, cover, temperature, impervious surfaces (roads and development), and topographic complexity (56% variation explained). We found that habitat permeability generally operates on fine scales, while topographic morphology and temperature-moisture regime operate across multiple scales, thus demonstrating the importance of cross-scale analysis for ecological interpretation. In a hierarchical analysis, we were able to explain more variation within genetic clusters as identified using Structure (a Bayesian algorithm) (74%; dispersal cover, growing-season precipitation, impervious surfaces) as opposed to between genetic clusters (45%; ridgelines, hot, dry slopes, length of hot season, and annual precipitation). Finally, the analytical methods we developed are powerful and can be applied to any species or system with appropriate landscape and genetic data.

Life-history evolution in range-shifting populations.

Abstract: Most evolutionary theory does not deal with populations expanding or contracting in space. Invasive species, climate change, epidemics, and the breakdown of dispersal barriers, however, all create populations in this kind of spatial disequilibrium. Importantly, spatial disequilibrium can have important ecological and evolutionary outcomes. During continuous range expansion, for example, populations on the expanding front experience novel evolutionary pressures because frontal populations are assorted by dispersal ability and have a lower density of conspecifics than do core populations. These conditions favor the evolution of traits that increase rates of dispersal and reproduction. Additionally, lowered density on the expanding front eventually frees populations on the expanding edge from specialist, coevolved enemies, permitting higher investment into traits associated with dispersal and reproduction rather than defense against pathogens. As a result, the process of range expansion drives rapid life-history evolution, and this seems to occur despite ongoing serial founder events that have complex effects on genetic diversity at the expanding front. Traits evolving on the expanding edge are smeared across the landscape as the front moves through, leaving an ephemeral signature of range expansion in the life-history traits of a species across its newly colonized range. Recent studies suggest that such nonequilibrium processes during recent population history may have contributed to many patterns usually ascribed to evolutionary forces acting in populations at spatial equilibrium.

Soil ecosystem functioning under climate change: plant species and community effects

Abstract: Feedbacks of terrestrial ecosystems to atmospheric and climate change depend on soil ecosystem dynamics. Soil ecosystems can directly and indirectly respond to climate change. For example, warming directly alters microbial communities by increasing their activity. Climate change may also alter plant community composition, thus indirectly altering the soil communities that depend on their inputs. To better understand how climate change may directly and indirectly alter soil ecosystem functioning, we investigated old-field plant community and soil ecosystem responses to single and combined effects of elevated [CO2], warming, and precipitation in Tennessee (USA). Specifically, we collected soils at the plot level (plant community soils) and beneath dominant plant species (plant-specific soils). We used microbial enzyme activities and soil nematodes as indicators for soil ecosystem functioning. Our study resulted in two main findings: (1) Overall, while there were some interactions, water, relative to increases in [CO2] and warming, had the largest impact on plant community composition, soil enzyme activity, and soil nematodes. Multiple climate-change factors can interact to shape ecosystems, but in our study, those interactions were largely driven by changes in water. (2) Indirect effects of climate change, via changes in plant communities, had a significant impact on soil ecosystem functioning, and this impact was not obvious when looking at plant community soils. Climate-change effects on enzyme activities and soil nematode abundance and community structure strongly differed between plant community soils and plant-specific soils, but also within plant-specific soils. These results indicate that accurate assessments of climate-change impacts on soil ecosystem functioning require incorporating the concurrent changes in plant function and plant community composition. Climate-change-induced shifts in plant community composition will likely modify or counteract the direct impact of atmospheric and climate change on soil ecosystem functioning, and hence, these indirect effects should be taken into account when predicting the manner in which global change will alter ecosystem functioning.

GAGES: A stream gage database for evaluating natural and altered flow conditions in the conterminous United States

Abstract: Stream flow is a controlling element in the ecology of rivers and streams. Knowledge of the natural flow regime facilitates the assessment of whether specific hydrologic attributes have been altered by humans in a particular stream and the establishment of specific goals for stream-flow restoration. Because most streams are ungaged or have been altered by human influences, characterizing the natural flow regime is often only possible by estimating flow characteristics based on nearby stream gages of reference quality, i.e., gaged locations that are least disturbed by human influences. The ability to evaluate natural stream flow, that which is not altered by human activities, would be enhanced by the existence of a nationally consistent and up-to-date database of gages in relatively undisturbed watersheds. As part of a national effort to characterize stream-flow effects on ecological condition, data for 6785 U.S. Geological Survey (USGS) stream gages and their upstream watersheds were compiled. The sites comprise all USGS stream gages in the conterminous United States with at least 20 years of complete-year flow record from 1950–2007, and for which watershed boundaries could reliably be delineated (median size = 578 km2). Several hundred watershed and site characteristics were calculated or compiled from national data sources, including environmental features (e.g., climate, geology, soils, topography) and anthropogenic influences (e.g., land use, roads, presence of dams, or canals). In addition, watersheds were assessed for their reference quality within nine broad regions for use in studies intended to characterize stream flows under conditions minimally influenced by human activities. Three primary criteria were used to assess reference quality: (1) a quantitative index of anthropogenic modification within the watershed based on GIS-derived variables, (2) visual inspection of every stream gage and drainage basin from recent high-resolution imagery and topographic maps, and (3) information about man-made influences from USGS Annual Water Data Reports. From the set of 6785 sites, we identified 1512 as reference-quality stream gages. All data derived for these watersheds as well as the reference condition evaluation are provided as an online data set termed GAGES (geospatial attributes of gages for evaluating stream flow). The complete data sets corresponding to abstracts published in the Data Papers section of the journal are published electronically in Ecological Archives at 〈http://esapubs.org/archive〉. (The accession number for each Data Paper is given directly beneath the title.)

Niche complementarity due to plasticity in resource use: plant partitioning of chemical N forms

Abstract: Niche complementarity, in which coexisting species use different forms of a resource, has been widely invoked to explain some of the most debated patterns in ecology, including maintenance of diversity and relationships between diversity and ecosystem function. However, classical models assume resource specialization in the form of distinct niches, which does not obviously apply to the broadly overlapping resource use in plant communities. Here we utilize an experimental framework based on competition theory to test whether plants partition resources via classical niche differentiation or via plasticity in resource use. We explore two alternatives: niche preemption, in which individuals respond to a superior competitor by switching to an alternative, less-used resource, and dominant plasticity, in which superior competitors exhibit high resource use plasticity and shift resource use depending on the competitive environment. We determined competitive ability by measuring growth responses with and without neighbors over a growing season and then used 15N tracer techniques to measure uptake of different nitrogen (N) forms in a field setting. We show that four alpine plant species of differing competitive abilities have statistically indistinguishable uptake patterns (nitrate > ammonium > glycine) in their fundamental niche (without competitors) but differ in whether they shift these uptake patterns in their realized niche (with competitors). Competitively superior species increased their uptake of the most available N form, ammonium, when in competition with the rarer, competitively inferior species. In contrast, the competitively inferior species did not alter its N uptake pattern in competition. The existence of plasticity in resource use among the dominant species provides a mechanism that helps to explain the manner by which plant species with broadly overlapping resource use might coexist.

Evidence of dispersal limitation in soil microorganisms: isolation reduces species richness on mycorrhizal tree islands.

Abstract: Dispersal limitation plays an important role in a number of equilibrium and nonequilibrium theories about community ecology. In this study we use the framework of island biogeography to look for evidence of dispersal limitation in ectomycorrhizal fungal assemblages on ''tree islands,'' patches of host trees located in a non-host vegetation matrix. Because of the potentially strong effects of island area on species richness and immigration, we chose to control island size by sampling tree islands consisting of a single host individual. Richness on tree islands was high, with estimates ranging up to 42 species of ectomycorrhizal fungi associating with a single host individual. Species richness decreased significantly with increasing isolation of tree islands, with our regression predicting a 50% decrease in species richness when tree islands are located distances of ;1 km from large patches of contiguous forests. Despite the fact that fungal fruit bodies produce large numbers of spores with high potential for long-distance travel, these results suggest that dispersal limitation is significant in ectomycorrhizal assemblages. There were no discernible effects of isolation or environment on the species identity of tree island fungal colonists. In contrast to the highly predictable patterns of tree island colonization we observed in a previous study on early successional forests, we suggest that over longer time periods the community assembly process becomes more dominated by stochastic immigration and local extinction events.

Winter climate change: a critical factor for temperate vegetation performance.

Abstract: Winter ecological processes are important drivers of vegetation and ecosystem functioning in temperate ecosystems. There, winter conditions are subject to rapid climate change. The potential loss of a longer-lasting snow cover with implications to other plant-related climate parameters and overwintering strategies make the temperate zone particularly vulnerable to winter climate change. A formalized literature search in the ISI Web of Science shows that plant related research on the effects of winter climate change is generally underrepresented. Temperate regions in particular are rarely studied in this respect, although the few existing studies imply strong effects of winter climate change on species ranges, species compositions, phenology, or frost injury. The generally positive effect of warming on plant survival and production may be counteracted by effects such as an increased frost injury of roots and shoots, an increased insect pest risk, or a disrupted synchrony between plants and pollinators. Based on the literature study, gaps in current knowledge are discussed. Understanding the relative effects of interacting climate parameters, as well as a stronger consideration of short-term events and variability of climatic conditions is urgent. With respect to plant response, it would be particularly worthwhile to account for hidden players such as pathogens, pollinators, herbivores, or fungal partners in mycorrhization.

Soil biota community structure and abundance under agricultural intensification and extensification.

Abstract: Understanding the impacts of agricultural intensification and extensification on soil biota communities is useful in order to preserve and restore biological diversity in agricultural soils and enhance the role of soil biota in agroecosystem functioning. Over four consecutive years, we investigated the effects of agricultural intensification and extensification (including conversion of grassland to arable land and vice versa, increased and decreased levels of mineral fertilization, and monoculture compared to crop rotation) on major soil biota group abundances and functional diversity. We integrated and compared effects across taxonomic levels to identify sensitive species groups. Conversion of grassland to arable land negatively affected both abundances and functional diversity of soil biota. Further intensification of the cropping system by increased fertilization and reduced crop diversity exerted smaller and differential effects on different soil biota groups. Agricultural intensification affected abundances of taxonomic groups with larger body size (earthworms, enchytraeids, microarthropods, and nematodes) more negatively than smaller-sized taxonomic groups (protozoans, bacteria, and fungi). Also functional group diversity and composition were more negatively affected in larger-sized soil biota (earthworms, predatory mites) than in smaller-sized soil biota (nematodes). Furthermore, larger soil biota appeared to be primarily affected by short-term consequences of conversion (disturbance, loss of habitat), whereas smaller soil biota were predominantly affected by long-term consequences (probably loss of organic matter). Reestablishment of grassland resulted in increased abundances of soil biota groups, but since not all groups increased in the same measure, the community structure was not completely restored. We concluded that larger-sized soil biota are more sensitive to agricultural intensification than smaller-sized soil biota. Furthermore, since larger-sized soil biota groups had lower taxonomic richness, we suggest that agricultural intensification exerts strongest effects on species-poor soil biota groups, thus supporting the hypothesis that biodiversity has an "insurance" function. As soil biota play an important role in agroecosystem functioning, altered soil biota abundances and functional group composition under agricultural intensification are likely to affect the functioning of the agroecosystem.

Global patterns in predator–prey size relationships reveal size dependency of trophic transfer efficiency

Abstract: Predator-prey body size relationships influence food chain length, trophic structure, transfer efficiency, interaction strength, and the bioaccumulation of contaminants. Improved quantification of these relationships and their response to the environment is needed to parameterize food web models and describe food web structure and function. A compiled data set comprising 29582 records of individual prey eaten at 21 locations by individual predators that spanned 10 orders of magnitude in mass and lived in marine environments ranging from the poles to the tropics was used to investigate the influence of predator size and environment on predator and prey size relationships. Linear mixed effects models demonstrated that predator-prey mass ratios (PPMR) increased with predator mass. The amount of the increase varied among locations and predator species and individuals but was not significantly influenced by temperature, latitude, depth, or primary production. Increases in PPMR with predator mass implied nonlinear relationships between log body mass and trophic level and reductions in transfer efficiency with increasing body size. The results suggest that very general rules determine dominant trends in PPMR in diverse marine ecosystems, leading to the ubiquity of size-based trophic structuring and the consistency of observed relationships between the relative abundance of individuals and their body size.

Are wolves saving Yellowstone's aspen? A landscape-level test of a behaviorally mediated trophic cascade

Abstract: Behaviorally mediated trophic cascades (BMTCs) occur when the fear of predation among herbivores enhances plant productivity. Based primarily on systems involving small-bodied predators, BMTCs have been proposed as both strong and ubiquitous in natural ecosystems. Recently, however, synthetic work has suggested that the existence of BMTCs may be mediated by predator hunting mode, whereby passive (sit-and-wait) predators have much stronger effects than active (coursing) predators. One BMTC that has been proposed for a wide-ranging active predator system involves the reintroduction of wolves (Canis lupus) to Yellowstone National Park, USA, which is thought to be leading to a recovery of trembling aspen (Populus tremuloides) by causing elk (Cervus elaphus) to avoid foraging in risky areas. Although this BMTC has been generally accepted and highly popularized, it has never been adequately tested. We assessed whether wolves influence aspen by obtaining detailed demographic data on aspen stands using tree rings and by monitoring browsing levels in experimental elk exclosures arrayed across a gradient of predation risk for three years. Our study demonstrates that the historical failure of aspen to regenerate varied widely among stands (last recruitment year ranged from 1892 to 1956), and our data do not indicate an abrupt cessation of recruitment. This pattern of recruitment failure appears more consistent with a gradual increase in elk numbers rather than a rapid behavioral shift in elk foraging following wolf extirpation. In addition, our estimates of relative survivorship of young browsable aspen indicate that aspen are not currently recovering in Yellowstone, even in the presence of a large wolf population. Finally, in an experimental test of the BMTC hypothesis we found that the impacts of elk browsing on aspen demography are not diminished in sites where elk are at higher risk of predation by wolves. These findings suggest the need to further evaluate how trophic cascades are mediated by predator-prey life history and ecological context.

Modeling species co‐occurrence by multivariate logistic regression generates new hypotheses on fungal interactions

Abstract: Signals of species interactions can be inferred from survey data by asking if some species occur more or less often together than what would be expected by random, or more generally, if any structural aspect of the community deviates from that expected from a set of independent species. However, a positive (or negative) association between two species does not necessarily signify a direct or indirect interaction, as it can result simply from the species having similar (or dissimilar) habitat requirements. We show how these two factors can be separated by multivariate logistic regression, with the regression part accounting for species-specific habitat requirements, and a correlation matrix for the positive or negative residual associations. We parameterize the model using Bayesian inference with data on 22 species of wood-decaying fungi acquired in 14 dissimilar forest sites. Our analyses reveal that some of the species commonly found to occur together in the same logs are likely to do so merely by similar habitat requirements, whereas other species combinations are systematically either over- or underrepresented also or only after accounting for the habitat requirements. We use our results to derive hypotheses on species interactions that can be tested in future experimental work.

Squeezed at the top: Interspecific aggression may constrain elevational ranges in tropical birds.

Abstract: Tropical montane species are characterized by narrow elevational distributions. Recent perspectives on mechanisms maintaining these restricted distributions have emphasized abiotic processes, but biotic processes may also play a role in their establishment or maintenance. One historically popular hypothesis, especially for birds, is that interspecific competition constrains ranges of closely related species that "replace" each other along elevational gradients. Supporting evidence, however, is based on patterns of occurrence and does not reveal potential mechanisms. We experimentally tested a prediction of this hypothesis in two genera of tropical songbirds, Catharus (Turdidae) and Henicorhina (Troglodytidae), in which species have nonoverlapping elevational distributions. Using heterospecific playback trials, we found that individuals at replacement zones showed aggressive territorial behavior in response to songs of congeners. As distance from replacement zones increased, aggression toward congener song decreased, suggesting a learned component to interspecific aggression. Additionally, aggressive responses in Catharus were asymmetric, indicating interspecific dominance. These results provide experimental evidence consistent with the hypothesis that interspecific competitive interactions restrict ranges of Neotropical birds. Our results also underscore the need to consider biotic processes, such as competition, when predicting how species' ranges will shift with climate change. Asymmetric aggression could be particularly important. For example, if warming in montane landscapes allows upslope range expansion by dominant competitors, then high-elevation subordinate species could be forced into progressively smaller mountaintop habitats, jeopardizing viability of their populations.

Climate change threatens polar bear populations: a stochastic demographic analysis

Abstract: The polar bear (Ursus maritimus) depends on sea ice for feeding, breeding, and movement. Significant reductions in Arctic sea ice are forecast to continue because of climate warming. We evaluated the impacts of climate change on polar bears in the southern Beaufort Sea by means of a demographic analysis, combining deterministic, stochastic, environment- dependent matrix population models with forecasts of future sea ice conditions from IPCC general circulation models (GCMs). The matrix population models classified individuals by age and breeding status; mothers and dependent cubs were treated as units. Parameter estimates were obtained from a capture-recapture study conducted from 2001 to 2006. Candidate statistical models allowed vital rates to vary with time and as functions of a sea ice covariate. Model averaging was used to produce the vital rate estimates, and a parametric bootstrap procedure was used to quantify model selection and parameter estimation uncertainty. Deterministic models projected population growth in years with more extensive ice coverage (2001-2003) and population decline in years with less ice coverage (2004-2005). LTRE (life table response experiment) analysis showed that the reduction in k in years with low sea ice was due primarily to reduced adult female survival, and secondarily to reduced breeding. A stochastic model with two environmental states, good and poor sea ice conditions, projected a declining stochastic growth rate, log ks, as the frequency of poor ice years increased. The observed frequency of poor ice years since 1979 would imply log ks ' � 0.01, which agrees with available (albeit crude) observations of population size. The stochastic model was linked to a set of 10 GCMs compiled by the IPCC; the models were chosen for their ability to reproduce historical observations of sea ice and were forced with ''business as usual'' (A1B) greenhouse gas emissions. The resulting stochastic population projections showed drastic declines in the polar bear population by the end of the 21st century. These projections were instrumental in the decision to list the polar bear as a threatened species under the U.S. Endangered Species Act.

Methane flux in non-wetland soils in response to nitrogen addition: a meta-analysis.

Abstract: The controls on methane (CH4) flux into and out of soils are not well understood Environmental variables including temperature, precipitation, and nitrogen (N) status can have strong effects on the magnitude and direction (eg, uptake vs release) of CH4 flux To better understand the interactions between CH4-cycling microorganisms and N in the non-wetland soil system, a meta-analysis was performed on published literature comparing CH4 flux in N amended and matched control plots An appropriate study index was developed for this purpose It was found that smaller amounts of N tended to stimulate CH4 uptake while larger amounts tended to inhibit uptake by the soil When all other variables were accounted for, the switch occurred at 100 kg N x ha(-1) x yr(-1) Managed land and land with a longer duration of fertilization showed greater inhibition of CH4 uptake with added N These results support the hypotheses that large amounts of available N can inhibit methanotrophy, but also that methanotrophs in upland soils can be N limited in their consumption of CH4 from the atmosphere There were interactions between other variables and N addition on the CH4 flux response: lower temperature and, to a lesser extent, higher precipitation magnified the inhibition of CH4 uptake due to N addition Several mechanisms that may cause these trends are discussed, but none could be conclusively supported with this approach Further controlled and in situ study should be undertaken to isolate the correct mechanism(s) responsible and to model upland CH4 flux

The ghosts of predators past: population cycles and the role of maternal programming under fluctuating predation risk

Abstract: Maternal effects may be a major factor influencing the demography of populations. In mammals, the transmission of stress hormones between mother and offspring may play an important role in these effects. Laboratory studies have shown that stressors during pregnancy and lactation result in lifelong programming of the offspring phenotype. However, the relevance of these studies to free-living mammals is unclear. The 10-year snowshoe hare (Lepus americanus) cycle is intimately linked to fluctuating predation pressure and predation risk. The enigma of these cycles is the lack of population growth following the decline phase, when the predators have virtually all disappeared and the food supply is ample. We have shown that a predator-induced increase in maternal stress hormone levels resulted in a decline in reproduction. Here we examine population and hormone changes over a four-year period from the increase (2005) to the decline (2008). We report (1) that an index of maternal stress (fecal corticosteroid metabolite [FCM] concentrations) fluctuates in synchrony with predator density during the breeding season; (2) that maternal FCM levels are echoed in their offspring, and this occurs at a population-wide level; and (3) that higher maternal FCM levels at birth are correlated with an increased responsiveness of the hypothalamic-pituitary-adrenal (HPA) axis in their progeny. Our results show an intergenerational inheritance of stress hormones in a free-ranging population of mammals. We propose that the lack of recovery of reproductive rates during the early low phase of the hare cycle may be the result of the impacts of intergenerational, maternally inherited stress hormones caused by high predation risk during the decline phase.

Meta-analyses and mega-mistakes: calling time on meta-analysis of the species richness-productivity relationship.

Abstract: The form of the species richness-productivity relationship (SRPR) is both theoretically important and contentious. In an effort to distill general patterns, ecologists have undertaken meta-analyses, within which each SRPR data set is first classified into one of five alternative forms: positive, humped (unimodal), negative, U-shaped (unimodal), and no relationship. Herein, I first provide a critique of this approach, based on 68 plant data sets/ studies used in three meta-analyses published in Ecology. The meta-analyses are shown to have resulted in highly divergent outcomes, inconsistent and often highly inappropriate classification of data sets, and the introduction and multiplication of errors from one meta-analysis to the next. I therefore call on the ecological community at large to adopt a far more rigorous and critical attitude to the use of meta-analysis. Second, I develop the argument that the literature on the SRPR continues to be bedeviled by a common failing to appreciate the fundamental importance of the scale of analysis, beginning with the confusion evident between concepts of grain, focus, and extent. I postulate that variation in the form of the SRPR at fine scales of analysis owes much to artifacts of the sampling regime adopted. An improved understanding may emerge from combining sampling theory with an understanding of the factors controlling the form of species abundance distributions and species accumulation curves.

A consumer–resource approach to the density-dependent population dynamics of mutualism

Abstract: Like predation and competition, mutualism is now recognized as a consumer- resource (C-R) interaction, including, in particular, bi-directional (e.g., coral, plant- mycorrhizae) and uni-directional (e.g., ant-plant defense, plant-pollinator) C-R mutualisms. Here, we develop general theory for the density-dependent population dynamics of mutualism based on the C-R mechanism of interspecific interaction. To test the influence of C-R interactions on the dynamics and stability of bi- and uni-directional C-R mutualisms, we developed simple models that link consumer functional response of one mutualistic species with the resources supplied by another. Phase-plane analyses show that the ecological dynamics of C-R mutualisms are stable in general. Most transient behavior leads to an equilibrium of mutualistic coexistence, at which both species densities are greater than in the absence of interactions. However, due to the basic nature of C-R interactions, certain density- dependent conditions can lead to C-R dynamics characteristic of predator-prey interactions, in which one species overexploits and causes the other to go extinct. Consistent with empirical phenomena, these results suggest that the C-R interaction can provide a broad mechanism for understanding density-dependent population dynamics of mutualism. By unifying predation, competition, and mutualism under the common ecological framework of consumer-resource theory, we may also gain a better understanding of the universal features of interspecific interactions in general.

On the evidence for species coexistence: a critique of the coexistence program

Abstract: A major challenge in ecology is to understand how the millions of species on Earth are organized into biological communities. Mechanisms promoting coexistence are one such class of organizing processes, which allow multiple species to persist in the same trophic level of a given web of species interactions. If some mechanism promotes the coexistence of two or more species, each species must be able to increase when it is rare and the others are at their typical abundances; this invasibility criterion is fundamental evidence for species coexistence regardless of the mechanism. In an attempt to evaluate the level of empirical support for coexistence mechanisms in nature, we surveyed the literature for empirical studies of coexistence at a local scale (i.e., species found living together in one place) to determine whether these studies satisfied the invasibility criterion. In our survey, only seven of 323 studies that drew conclusions about species coexistence evaluated invasibility in some way in either observational or experimental studies. In addition, only three other studies evaluated necessary but not sufficient conditions for invasibility (i.e., negative density dependence and a trade-off in performance that influences population regulation). These results indicate that, while species coexistence is a prevalent assumption for why species are able to live together in one place, critical empirical tests of this fundamental assumption of community structure are rarely performed. These tests are central to developing a more robust understanding of the relative contributions of both deterministic and stochastic processes structuring biological communities.

Photoacclimatization by the coral Montastraea cavernosa in the mesophotic zone: light, food, and genetics

Abstract: Most studies on coral reefs have focused on shallow reef (<30 m) systems due to the technical limitations of conducting scientific diving deeper than 30 m. Compared to their shallow-water counterparts, these mesophotic coral reefs (30-150 m) are understudied, which has slowed our broader understanding of the biodiversity, ecology, and connectivity of shallow and deep coral reef communities. We know that the light environment is an important component of the productivity, physiology, and ecology of corals, and it restricts the distribution of most species of coral to depths of 60 m or less. In the Bahamas, the coral Montastraea cavernosa has a wide depth distribution, and it is one of the most numerous corals at mesophotic depths. Using a range of optical, physiological, and biochemical approaches, the relative dependence on autotrophy vs. heterotrophy was assessed for this coral from 3 to 91 m. These measurements show that the quantum yield of PSII fluorescence increases significantly with depth for M. cavernosa while gross primary productivity decreases with depth. Both morphological and physiological photoacclimatization occurs to a depth of 91 m, and stable isotope data of the host tissues, symbionts, and skeleton reveal a marked decrease in productivity and a sharp transition to heterotrophy between 45 and 61 m. Below these depths, significant changes in the genetic composition of the zooxanthellae community, including genotypes not previously observed, occur and suggest that there is strong selection for zooxanthellae that are suited for survival in the light-limited environment where mesophotic M. cavernosa are occurring.

Divergent composition but similar function of soil food webs of individual plants: plant species and community effects

Abstract: Soils are extremely rich in biodiversity, and soil organisms play pivotal roles in supporting terrestrial life, but the role that individual plants and plant communities play in influencing the diversity and functioning of soil food webs remains highly debated Plants, as primary producers and providers of resources to the soil food web, are of vital importance for the composition, structure, and functioning of soil communities However, whether natural soil food webs that are completely open to immigration and emigration differ underneath individual plants remains unknown In a biodiversity restoration experiment we first compared the soil nematode communities of 228 individual plants belonging to eight herbaceous species We included grass, leguminous, and non-leguminous species Each individual plant grew intermingled with other species, but all plant species had a different nematode community Moreover, nematode communities were more similar when plant individuals were growing in the same as compared to different plant communities, and these effects were most apparent for the groups of bacterivorous, carnivorous, and omnivorous nematodes Subsequently, we analyzed the composition, structure, and functioning of the complete soil food webs of 58 individual plants, belonging to two of the plant species, Lotus corniculatus (Fabaceae) and Plantago lanceolata (Plantaginaceae) We isolated and identified more than 150 taxa/groups of soil organisms The soil community composition and structure of the entire food webs were influenced both by the species identity of the plant individual and the surrounding plant community Unexpectedly, plant identity had the strongest effects on decomposing soil organisms, widely believed to be generalist feeders In contrast, quantitative food web modeling showed that the composition of the plant community influenced nitrogen mineralization under individual plants, but that plant species identity did not affect nitrogen or carbon mineralization or food web stability Hence, the composition and structure of entire soil food webs vary at the scale of individual plants and are strongly influenced by the species identity of the plant However, the ecosystem functions these food webs provide are determined by the identity of the entire plant community