Showing papers in "Ecology in 2018"

Species co-occurrence networks: Can they reveal trophic and non-trophic interactions in ecological communities?

Abstract: Co-occurrence methods are increasingly utilized in ecology to infer networks of species interactions where detailed knowledge based on empirical studies is difficult to obtain. Their use is particularly common, but not restricted to, microbial networks constructed from metagenomic analyses. In this study, we test the efficacy of this procedure by comparing an inferred network constructed using spatially intensive co-occurrence data from the rocky intertidal zone in central Chile to a well-resolved, empirically based, species interaction network from the same region. We evaluated the overlap in the information provided by each network and the extent to which there is a bias for co-occurrence data to better detect known trophic or non-trophic, positive or negative interactions. We found a poor correspondence between the co-occurrence network and the known species interactions with overall sensitivity (probability of true link detection) equal to 0.469, and specificity (true non-interaction) equal to 0.527. The ability to detect interactions varied with interaction type. Positive non-trophic interactions such as commensalism and facilitation were detected at the highest rates. These results demonstrate that co-occurrence networks do not represent classical ecological networks in which interactions are defined by direct observations or experimental manipulations. Co-occurrence networks provide information about the joint spatial effects of environmental conditions, recruitment, and, to some extent, biotic interactions, and among the latter, they tend to better detect niche-expanding positive non-trophic interactions. Detection of links (sensitivity or specificity) was not higher for well-known intertidal keystone species than for the rest of consumers in the community. Thus, as observed in previous empirical and theoretical studies, patterns of interactions in co-occurrence networks must be interpreted with caution, especially when extending interaction-based ecological theory to interpret network variability and stability. Co-occurrence networks may be particularly valuable for analysis of community dynamics that blends interactions and environment, rather than pairwise interactions alone.

PHYLACINE 1.2: The Phylogenetic Atlas of Mammal Macroecology

Abstract: Data needed for macroecological analyses are difficult to compile and often hidden away in supplementary material under non-standardized formats. Phylogenies, range data, and trait data often use conflicting taxonomies and require ad hoc decisions to synonymize species or fill in large amounts of missing data. Furthermore, most available data sets ignore the large impact that humans have had on species ranges and diversity. Ignoring these impacts can lead to drastic differences in diversity patterns and estimates of the strength of biological rules. To help overcome these issues, we assembled PHYLACINE, The Phylogenetic Atlas of Mammal Macroecology. This taxonomically integrated platform contains phylogenies, range maps, trait data, and threat status for all 5,831 known mammal species that lived since the last interglacial (~130,000 years ago until present). PHYLACINE is ready to use directly, as all taxonomy and metadata are consistent across the different types of data, and files are provided in easy-to-use formats. The atlas includes both maps of current species ranges and present natural ranges, which represent estimates of where species would live without anthropogenic pressures. Trait data include body mass and coarse measures of life habit and diet. Data gaps have been minimized through extensive literature searches and clearly labelled imputation of missing values. The PHYLACINE database will be archived here as well as hosted online so that users may easily contribute updates and corrections to continually improve the data. This database will be useful to any researcher who wishes to investigate large-scale ecological patterns. Previous versions of the database have already provided valuable information and have, for instance, shown that megafauna extinctions caused substantial changes in vegetation structure and nutrient transfer patterns across the globe.

Microbes Follow Humboldt: Temperature Drives Plant and Soil Microbial Diversity Patterns from the Amazon to the Andes

Abstract: More than 200 years ago, Alexander von Humboldt reported that tropical plant species richness decreased with increasing elevation and decreasing temperature. Surprisingly, coordinated patterns in plant, bacterial, and fungal diversity on tropical mountains have not yet been observed, despite the central role of soil microorganisms in terrestrial biogeochemistry and ecology. We studied an Andean transect traversing 3.5 km in elevation to test whether the species diversity and composition of tropical forest plants, soil bacteria, and fungi follow similar biogeographical patterns with shared environmental drivers. We found coordinated changes with elevation in all three groups: species richness declined as elevation increased, and the compositional dissimilarity among communities increased with increased separation in elevation, although changes in plant diversity were larger than in bacteria and fungi. Temperature was the dominant driver of these diversity gradients, with weak influences of edaphic properties, including soil pH. The gradients in microbial diversity were strongly correlated with the activities of enzymes involved in organic matter cycling, and were accompanied by a transition in microbial traits towards slower-growing, oligotrophic taxa at higher elevations. We provide the first evidence of coordinated temperature-driven patterns in the diversity and distribution of three major biotic groups in tropical ecosystems: soil bacteria, fungi, and plants. These findings suggest that interrelated and fundamental patterns of plant and microbial communities with shared environmental drivers occur across landscape scales. These patterns are revealed where soil pH is relatively constant, and have implications for tropical forest communities under future climate change.

Ecological drivers of soil microbial diversity and soil biological networks in the Southern Hemisphere.

Abstract: The ecological drivers of soil biodiversity in the Southern Hemisphere remain underexplored. Here, in a continental survey comprising 647 sites, across 58 degrees of latitude between tropical Australia and Antarctica, we evaluated the major ecological patterns in soil biodiversity and relative abundance of ecological clusters within a co-occurrence network of soil bacteria, archaea and eukaryotes. Six major ecological clusters (modules) of co-occurring soil taxa were identified. These clusters exhibited strong shifts in their relative abundances with increasing distance from the equator. Temperature was the major environmental driver of the relative abundance of ecological clusters when Australia and Antarctica are analyzed together. Temperature, aridity, soil properties and vegetation types were the major drivers of the relative abundance of different ecological clusters within Australia. Our data supports significant reductions in the diversity of bacteria, archaea and eukaryotes in Antarctica vs. Australia linked to strong reductions in temperature. However, we only detected small latitudinal variations in soil biodiversity within Australia. Different environmental drivers regulate the diversity of soil archaea (temperature and soil carbon), bacteria (aridity, vegetation attributes and pH) and eukaryotes (vegetation type and soil carbon) across Australia. Together, our findings provide new insights into the mechanisms driving soil biodiversity in the Southern Hemisphere.

On the relationship between phylogenetic diversity and trait diversity

Abstract: Niche differences are key to understanding the distribution and structure of biodiversity. To examine niche differences, we must first characterize how species occupy niche space, and two approaches are commonly used in the ecological literature. The first uses species traits to estimate multivariate trait space (so-called functional trait diversity, FD); the second quantifies the amount of time or evolutionary history captured by a group of species (phylogenetic diversity, PD). It is often-but controversially-assumed that these putative measures of niche space are at a minimum correlated and perhaps redundant, since more evolutionary time allows for greater accumulation of trait changes. This theoretical expectation remains surprisingly poorly evaluated, particularly in the context of multivariate measures of trait diversity. We evaluated the relationship between phylogenetic diversity and trait diversity using analytical and simulation-based methods across common models of trait evolution. We show that PD correlates with FD increasingly strongly as more traits are included in the FD measure. Our results indicate that phylogenetic diversity can be a useful surrogate for high-dimensional trait diversity, but we also show that the correlation weakens when the underlying process of trait evolution includes variation in rate and optima.

Competition for nutrients and light: testing advances in resource competition with a natural phytoplankton community

Abstract: A key challenge in ecology is to understand how nutrients and light affect the biodiversity and community structure of phytoplankton and plant communities. According to resource competition models, ratios of limiting nutrients are major determinants of species composition. At high nutrient levels, however, species interactions may shift to competition for light, which might make nutrient ratios less relevant. The "nutrient-load hypothesis" merges these two perspectives, by extending the classic model of competition for two nutrients to include competition for light. Here, we test five key predictions of the nutrient-load hypothesis using multispecies competition experiments. A marine phytoplankton community sampled from the North Sea was inoculated in laboratory chemostats provided with different nitrogen (N) and phosphorus (P) loads to induce either single resource limitation or co-limitation of N, P, and light. Four of the five predictions were validated by the experiments. In particular, different resource limitations favored the dominance of different species. Increasing nutrient loads caused changes in phytoplankton species composition, even if the N:P ratio of the nutrient loads remained constant, by shifting the species interactions from competition for nutrients to competition for light. In all treatments, small species became dominant whereas larger species were competitively excluded, supporting the common view that small cell size provides a competitive advantage under resource-limited conditions. Contrary to expectation, all treatments led to coexistence of diatoms, cyanobacteria and green algae, resulting in a higher diversity of species than predicted by theory. Because the coexisting species comprised three phyla with different photosynthetic pigments, we speculate that niche differentiation in the light spectrum might play a role. Our results show that mechanistic resource competition models that integrate nutrient-based and light-based approaches provide an important step forward to understand and predict how changing nutrient loads affect community composition.

Identifiability in N-mixture models: a large-scale screening test with bird data

Abstract: Binomial N-mixture models have proven very useful in ecology, conservation, and monitoring: they allow estimation and modeling of abundance separately from detection probability using simple counts. Recently, doubts about parameter identifiability have been voiced. I conducted a large-scale screening test with 137 bird data sets from 2,037 sites. I found virtually no identifiability problems for Poisson and zero-inflated Poisson (ZIP) binomial N-mixture models, but negative-binomial (NB) models had problems in 25% of all data sets. The corresponding multinomial N-mixture models had no problems. Parameter estimates under Poisson and ZIP binomial and multinomial N-mixture models were extremely similar. Identifiability problems became a little more frequent with smaller sample sizes (267 and 50 sites), but were unaffected by whether the models did or did not include covariates. Hence, binomial N-mixture model parameters with Poisson and ZIP mixtures typically appeared identifiable. In contrast, NB mixtures were often unidentifiable, which is worrying since these were often selected by Akaike's information criterion. Identifiability of binomial N-mixture models should always be checked. If problems are found, simpler models, integrated models that combine different observation models or the use of external information via informative priors or penalized likelihoods, may help.

On the robustness of N-mixture models

Abstract: N-mixture models provide an appealing alternative to mark-recapture models, in that they allow for estimation of detection probability and population size from count data, without requiring that individual animals be identified. There is, however, a cost to using the N-mixture models: inference is very sensitive to the model's assumptions. We consider the effects of three violations of assumptions that might reasonably be expected in practice: double counting, unmodeled variation in population size over time, and unmodeled variation in detection probability over time. These three examples show that small violations of assumptions can lead to large biases in estimation. The violations of assumptions we consider are not only small qualitatively, but are also small in the sense that they are unlikely to be detected using goodness-of-fit tests. In cases where reliable estimates of population size are needed, we encourage investigators to allocate resources to acquiring additional data, such as recaptures of marked individuals, for estimation of detection probabilities.

Moisture availability limits subalpine tree establishment.

Abstract: In the absence of broad-scale disturbance, many temperate coniferous forests experience successful seedling establishment only when abundant seed production coincides with favorable climate. Identifying the frequency of past establishment events and the climate conditions favorable for seedling establishment is essential to understanding how climate warming could affect the frequency of future tree establishment events and therefore future forest composition or even persistence of a forest cover. In the southern Rocky Mountains, USA, research on the sensitivity of establishment of Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa)-two widely distributed, co-occurring conifers in North America-to climate variability has focused on the alpine treeline ecotone, leaving uncertainty about the sensitivity of these species across much of their elevation distribution. We compared annual germination dates for >450 Engelmann spruce and >500 subalpine fir seedlings collected across a complex topographic-moisture gradient to climate variability in the Colorado Front Range. We found that Engelmann spruce and subalpine fir established episodically with strong synchrony in establishment events across the study area. Broad-scale establishment events occurred in years of high soil moisture availability, which were characterized by above-average snowpack and/or cool and wet summer climatic conditions. In the recent half of the study period (1975-2010), a decrease in the number of fir and spruce establishment events across their distribution coincided with declining snowpack and a multi-decadal trend of rising summer temperature and increasing moisture deficits. Counter to expected and observed increases in tree establishment with climate warming in maritime subalpine forests, our results show that recruitment declines will likely occur across the core of moisture-limited subalpine tree ranges as warming drives increased moisture deficits.

Tick-borne disease risk in a forest food web.

Abstract: Changes to the community ecology of hosts for zoonotic pathogens, particularly rodents, are likely to influence the emergence and prevalence of zoonotic diseases worldwide. However, the complex interactions between abiotic factors, pathogens, vectors, hosts, and both food resources and predators of hosts are difficult to disentangle. Here we (1) use 19 yr of data from six large field plots in southeastern New York to compare the effects of hypothesized drivers of interannual variation in Lyme disease risk, including the abundance of acorns, rodents, and deer, as well as a series of climate variables; and (2) employ landscape epidemiology to explore how variation in predator community structure and forest cover influences spatial variation in the infection prevalence of ticks for the Lyme disease bacterium, Borrelia burgdorferi, and two other important tick-borne pathogens, Anaplasma phagocytophilum and Babesia microti. Acorn-driven increases in the abundance of mice were correlated with a lagged increase in the abundance of questing nymph-stage Ixodes scapularis ticks infected with Lyme disease bacteria. Abundance of white-tailed deer 2 yr prior also correlated with increased density of infected nymphal ticks, although the effect was weak. Density of rodents in the current year was a strong negative predictor of nymph density, apparently because high current abundance of these hosts can remove nymphs from the host-seeking population. Warm, dry spring or winter weather was associated with reduced density of infected nymphs. At the landscape scale, the presence of functionally diverse predator communities or of bobcats, the only obligate carnivore, was associated with reduced infection prevalence of I. scapularis nymphs with all three zoonotic pathogens. In the case of Lyme disease, infection prevalence increased where coyotes were present but smaller predators were displaced or otherwise absent. For all pathogens, infection prevalence was lowest when forest cover within a 1 km radius was high. Taken together, our results suggest that a food web perspective including bottom-up and top-down forcing is needed to understand drivers of tick-borne disease risk, a result that may also apply to other rodent-borne zoonoses. Prevention of exposure based on ecological indicators of heightened risk should help protect public health.

Plant responses to fertilization experiments in lowland, species-rich, tropical forests.

Abstract: We present a meta-analysis of plant responses to fertilization experiments conducted in lowland, species-rich, tropical forests. We also update a key result and present the first species-level analyses of tree growth rates for a 15-yr factorial nitrogen (N), phosphorus (P), and potassium (K) experiment conducted in central Panama. The update concerns community-level tree growth rates, which responded significantly to the addition of N and K together after 10 yr of fertilization but not after 15 yr. Our experimental soils are infertile for the region, and species whose regional distributions are strongly associated with low soil P availability dominate the local tree flora. Under these circumstances, we expect muted responses to fertilization, and we predicted species associated with low-P soils would respond most slowly. The data did not support this prediction, species-level tree growth responses to P addition were unrelated to species-level soil P associations. The meta-analysis demonstrated that nutrient limitation is widespread in lowland tropical forests and evaluated two directional hypotheses concerning plant responses to N addition and to P addition. The meta-analysis supported the hypothesis that tree (or biomass) growth rate responses to fertilization are weaker in old growth forests and stronger in secondary forests, where rapid biomass accumulation provides a nutrient sink. The meta-analysis found no support for the long-standing hypothesis that plant responses are stronger for P addition and weaker for N addition. We do not advocate discarding the latter hypothesis. There are only 14 fertilization experiments from lowland, species-rich, tropical forests, 13 of the 14 experiments added nutrients for five or fewer years, and responses vary widely among experiments. Potential fertilization responses should be muted when the species present are well adapted to nutrient-poor soils, as is the case in our experiment, and when pest pressure increases with fertilization, as it does in our experiment. The statistical power and especially the duration of fertilization experiments conducted in old growth, tropical forests might be insufficient to detect the slow, modest growth responses that are to be expected.

Optimizing the choice of a spatial weighting matrix in eigenvector‐based methods

Abstract: Eigenvector-mapping methods such as Moran's eigenvector maps (MEM) are derived from a spatial weighting matrix (SWM) that describes the relations among a set of sampled sites The specification of the SWM is a crucial step, but the SWM is generally chosen arbitrarily, regardless of the sampling design characteristics Here, we compare the statistical performances of different types of SWMs (distance-based or graph-based) in contrasted realistic simulation scenarios Then, we present an optimization method and evaluate its performances compared to the arbitrary choice of the most-widely used distance-based SWM Results showed that the distance-based SWMs generally had lower power and accuracy than other specifications, and strongly underestimated spatial signals The optimization method, using a correction procedure for multiple tests, had a correct type I error rate, and had higher power and accuracy than an arbitrary choice of the SWM Nevertheless, the power decreased when too many SWMs were compared, resulting in a trade-off between the gain of accuracy and the loss of power We advocate that future studies should optimize the choice of the SWM using a small set of appropriate candidates R functions to implement the optimization are available in the adespatial package and are detailed in a tutorial

Resource availability underlies the plant-fungal diversity relationship in a grassland ecosystem

Abstract: It is commonly assumed that microbial communities are structured by "bottom-up" ecological forces, although few experimental manipulations have rigorously tested the mechanisms by which resources structure soil communities. We investigated how plant substrate availability might structure fungal communities and belowground processes along an experimental plant richness gradient in a grassland ecosystem. We hypothesized that variation in total plant-derived substrate inputs, plant functional group diversity, as well as the relative abundance of C4 grasses and legumes would modulate fungal α- and β-diversity and their rates of soil carbon (C) and nitrogen (N) cycling. To test these predictions, we molecularly characterized fungal communities, as well as potential extracellular enzyme activity, net N mineralization, and soil organic matter respiration. We found higher fungal richness was associated with increasing aboveground plant biomass; whereas, fungal β-diversity was explained by contributions from C4 grass and legume relative dominance, plant functional group diversity, as well as plant biomass. Furthermore, aboveground plant biomass consistently shaped the richness and composition of individual fungal trophic modes (i.e., saprotrophs, symbiotrophs, pathotrophs). Finally, variation in extracellular enzyme activity, net N mineralization rates, and soil organic matter respiration was significantly explained by fungal β-diversity when fungi were functionally classified. Via changes in the supply and composition of organic substrates entering soil, our study demonstrates that changes in the plant species richness and functional composition collectively influence fungal communities and rates of soil C and N cycling.

Sampling scales define occupancy and underlying occupancy-abundance relationships in animals

Abstract: Occupancy-abundance (OA) relationships are a foundational ecological phenomenon and field of study, and occupancy models are increasingly used to track population trends and understand ecological interactions. However, these two fields of ecological inquiry remain largely isolated, despite growing appreciation of the importance of integration. For example, using occupancy models to infer trends in abundance is predicated on positive OA relationships. Many occupancy studies collect data that violate geographical closure assumptions due to the choice of sampling scales and application to mobile organisms, which may change how occupancy and abundance are related. Little research, however, has explored how different occupancy sampling designs affect OA relationships. We develop a conceptual framework for understanding how sampling scales affect the definition of occupancy for mobile organisms, which drives OA relationships. We explore how spatial and temporal sampling scales, and the choice of sampling unit (areal vs. point sampling), affect OA relationships. We develop predictions using simulations, and test them using empirical occupancy data from remote cameras on 11 medium-large mammals. Surprisingly, our simulations demonstrate that when using point sampling, OA relationships are unaffected by spatial sampling grain (i.e., cell size). In contrast, when using areal sampling (e.g., species atlas data), OA relationships are affected by spatial grain. Furthermore, OA relationships are also affected by temporal sampling scales, where the curvature of the OA relationship increases with temporal sampling duration. Our empirical results support these predictions, showing that at any given abundance, the spatial grain of point sampling does not affect occupancy estimates, but longer surveys do increase occupancy estimates. For rare species (low occupancy), estimates of occupancy will quickly increase with longer surveys, even while abundance remains constant. Our results also clearly demonstrate that occupancy for mobile species without geographical closure is not true occupancy. The independence of occupancy estimates from spatial sampling grain depends on the sampling unit. Point-sampling surveys can, however, provide unbiased estimates of occupancy for multiple species simultaneously, irrespective of home-range size. The use of occupancy for trend monitoring needs to explicitly articulate how the chosen sampling scales define occupancy and affect the occupancy-abundance relationship.

It takes a few to tango: changing climate and fire regimes can cause regeneration failure of two subalpine conifers.

Abstract: Environmental change is accelerating in the 21st century, but how multiple drivers may interact to alter forest resilience remains uncertain. In forests affected by large high-severity disturbances, tree regeneration is a resilience linchpin that shapes successional trajectories for decades. We modeled stands of two widespread western U.S. conifers, Douglas-fir (Pseudotsuga menziesii var. glauca), and lodgepole pine (Pinus contorta var. latifolia), in Yellowstone National Park (Wyoming, USA) to ask (1) What combinations of distance to seed source, fire return interval, and warming-drying conditions cause postfire tree-regeneration failure? (2) If postfire tree regeneration was successful, how does early tree density differ under future climate relative to historical climate? We conducted a stand-level (1 ha) factorial simulation experiment using the individual-based forest process model iLand to identify combinations of fire return interval (11-100 yr), distance to seed source (50-1,000 m), and climate (historical, mid-21st century, late-21st century) where trees failed to regenerate by 30-yr postfire. If regeneration was successful, we compared stand densities between climate periods. Simulated postfire regeneration were surprisingly resilient to changing climate and fire drivers. Douglas-fir regeneration failed more frequently (55%) than lodgepole pine (28% and 16% for non-serotinous and serotinous stands, respectively). Distance to seed source was an important driver of regeneration failure for Douglas-fir and non-serotinous lodgepole pine; regeneration never failed when stands were 50 m from a seed source and nearly always failed when stands were 1 km away. Regeneration of serotinous lodgepole pine only failed when fire return intervals were ≤20 yr and stands were far (1 km) from a seed source. Warming climate increased regeneration success for Douglas-fir but did not affect lodgepole pine. If regeneration was successful, postfire density varied with climate. Douglas-fir and serotinous lodgepole pine regeneration density both increased under 21st-century climate but in response to different climate variables (growing season length vs. cold limitation). Results suggest that, given a warmer future with larger and more frequent fires, a greater number of stands that fail to regenerate after fires combined with increasing density in stands where regeneration is successful could produce a more coarse-grained forest landscape.

Fundamental contradictions among observational and experimental estimates of non-trophic species interactions

Abstract: The difficulty of experimentally quantifying non-trophic species interactions has long troubled ecologists. Increasingly, a new application of the classic "checkerboard distribution" approach is used to infer interactions by examining the pairwise frequency at which species are found to spatially co-occur. However, the link between spatial associations, as estimated from observational co-occurrence, and species interactions has not been tested. Here we used nine common statistical methods to estimate associations from surveys of rocky intertidal communities in the Northeast Pacific Ocean. We compared those inferred associations with a new data set of experimentally determined net and direct species interactions. Although association methods generated networks with aggregate structure similar to previously published interaction networks, each method detected a different set of species associations from the same data set. Moreover, although association methods generally performed better than a random model, associations rarely matched empirical net or direct species interactions, with high rates of false positives and true positives, and many false negatives. Our findings cast doubt on studies that equate species co-occurrences to species interactions and highlight a persistent, unanswered question: how do we interpret spatial patterns in communities? We suggest future research directions to unify the observational and experimental study of species interactions, and discuss the need for community standards and best practices in association analysis.

Functional diversity positively affects prey suppression by invertebrate predators: a meta-analysis.

Abstract: The use of pesticides within agricultural ecosystems has led to wide concern regarding negative effects on the environment. One possible alternative is the use of predators of pest species that naturally occur within agricultural ecosystems. However, the mechanistic basis for how species can be manipulated in order to maximize pest control remains unclear. We carried out a meta-analysis of 51 studies that manipulated predator species richness in reference to suppression of herbivore prey to determine which components of predator diversity affect pest control. Overall, functional diversity (FD) based on predator's habitat domain, diet breadth and hunting strategy was ranked as the most important variable. Our analysis showed that increases in FD in polycultures led to greater prey suppression compared to both the mean of the component predator species, and the most effective predator species, in monocultures. Further analysis of individual traits indicated these effects are likely to be driven by broad niche differentiation and greater resource exploitation in functionally diverse predator communities. A decoupled measure of phylogenetic diversity, whereby the overlap in variation with FD was removed, was not found to be an important driver of prey suppression. Our results suggest that increasing FD in predatory invertebrates will help maximize pest control ecosystem services in agricultural ecosystems, with the potential to increase suppression above that of the most effective predator species.

Freshwater eutrophication drives sharp reductions in temporal beta diversity.

Abstract: Eutrophication has become one of the most widespread anthropogenic forces impacting freshwater biological diversity. One potentially important mechanism driving biodiversity changes in response to eutrophication is the alteration of seasonal patterns of succession, particularly among species with short, synchronous, life cycles. We tested the hypothesis that eutrophication reduces seasonally driven variation in species assemblages by focusing on an understudied aspect of biodiversity: temporal beta diversity (βt ). We estimated the effect of eutrophication on βt by sampling benthic macroinvertebrate assemblages bimonthly for two years across 35 streams spanning a steep gradient of total phosphorus (P) and benthic algal biomass (as chlorophyll a [chl a]). Two widely used metrics of β diversity both declined sharply in response to increasing P and chl a, regardless of covariates. The most parsimonious explanatory model for βt included an interaction between P and macroinvertebrate biomass, which revealed that βt was lower when macroinvertebrate biomass was relatively high. Macroinvertebrate biomass explained a greater amount of deviance in βt at lower to moderate concentrations of P, providing additional explanatory power where P concentration alone was unable to fully explain declines in βt . Chl a explained similar amounts of deviance in βt in comparison to the best P model, but only when temperature variability, which was positively related to βt , also was included in the model. Declines in βt suggest that nutrient enrichment decreases the competitive advantage that specialists gain by occupying particular temporal niches, which leads to assemblages dominated by generalists that exhibit little seasonal turnover. The collapse of seasonal variation in assemblage composition we observed in our study suggests that treating dynamic communities as static assemblages is a simplification that may fail to detect the full impact of anthropogenic stressors. Our results show that eutrophication leads to more temporally homogenous communities and therefore degrades a fundamental facet of biodiversity.

Climate sensitivity functions and net primary production: A framework for incorporating climate mean and variability.

Abstract: Understanding controls on net primary production (NPP) has been a long-standing goal in ecology. Climate is a well-known control on NPP, although the temporal differences among years within a site are often weaker than the spatial pattern of differences across sites. Climate sensitivity functions describe the relationship between an ecological response (e.g., NPP) and both the mean and variance of its climate driver (e.g., aridity index), providing a novel framework for understanding how climate trends in both mean and variance vary with NPP over time. Nonlinearities in these functions predict whether an increase in climate variance will have a positive effect (convex nonlinearity) or negative effect (concave nonlinearity) on NPP. The influence of climate variance may be particularly intense at ecosystem transition zones, if species reach physiological thresholds that create nonlinearities at these ecotones. Long-term data collected at the confluence of three dryland ecosystems in central New Mexico revealed that each ecosystem exhibited a unique climate sensitivity function that was consistent with long-term vegetation change occurring at their ecotones. Our analysis suggests that rising temperatures in drylands could alter the nonlinearities that determine the relative costs and benefits of variance in precipitation for primary production.

Topography and neighborhood crowding can interact to shape species growth and distribution in a diverse Amazonian forest.

Abstract: Abiotic constraints and biotic interactions act simultaneously to shape communities. However, these community assembly mechanisms are often studied independently, which can limit understanding of how they interact to affect species dynamics and distributions. We develop a hierarchical Bayesian neighborhood modeling approach to quantify the simultaneous effects of topography and crowding by neighbors on the growth of 124,704 individual stems 1cm DBH for 1,047 tropical tree species in a 25-ha mapped rainforest plot in Amazonian Ecuador. We build multi-level regression models to evaluate how four key functional traits (specific leaf area, maximum tree size, wood specific gravity and seed mass) mediate tree growth response to topography and neighborhood crowding. Tree growth is faster in valleys than on ridges and is reduced by neighborhood crowding. Topography and crowding interact to influence tree growth in similar to 10% of the species. Specific leaf area, maximum tree size and seed mass are associated with growth responses to topography, but not with responses to neighborhood crowding or with the interaction between topography and crowding. In sum, our study reveals that topography and neighborhood crowding each influence tree growth in tropical forests, but act largely independently in shaping species distributions. While traits were associated with species response to topography, their role in species response to neighborhood crowding was less clear, which suggests that trait effects on neighborhood dynamics may depend on the direction (negative/positive) and degree of symmetry of biotic interactions. Our study emphasizes the importance of simultaneously assessing the individual and interactive role of multiple mechanisms in shaping species dynamics in high diversity tropical systems.

Plant diversity and density predict belowground diversity and function in an early successional alpine ecosystem.

Abstract: Despite decades of interest, few studies have provided evidence supporting theoretical expectations for coupled relationships between aboveground and belowground diversity and ecosystem functioning in non-manipulated natural ecosystems We characterized plant species richness and density, soil bacterial, fungal and eukaryotic species richness and phylogenetic diversity (using 16S, ITS, and 18S gene sequencing), and ecosystem function (levels of soil C and N, and rates of microbial enzyme activities) along a natural gradient in plant richness and density in high-elevation, C-deficient soils to examine the coupling between above- and belowground systems Overall, we observed a strong positive relationship between aboveground (plant richness and density) and belowground (bacteria, fungi, and non-fungal eukaryotes) richness In addition to the correlations between plants and soil communities, C and N pools, and rates of enzyme activities increased as plant and soil communities became richer and more diverse Our results suggest that the theoretically expected positive correlation between above- and belowground communities does exist in natural systems, but may be undetectable in late successional ecosystems due to the buildup of legacy organic matter that results in extremely complex belowground communities In contrast, microbial communities in early successional systems, such as the system described here, are more directly dependent on contemporary inputs from plants and therefore are strongly correlated with plant diversity and density

Interaction frequency, network position, and the temporal persistence of interactions in a plant-pollinator network.

Abstract: Ecological interactions are highly dynamic in time and space. Previous studies of plant-animal mutualistic networks have shown that the occurrence of interactions varies substantially across years. We analyzed interannual variation of a quantitative mutualistic network, in which links are weighted by interaction frequency. The network was sampled over six consecutive years, representing one of the longest time series for a community-wide mutualistic network. We estimated the interannual similarity in interactions and assessed the determinants of their persistence. The occurrence of interactions varied greatly among years, with most interactions seen in only one year (64%) and few (20%) in more than two years. This variation was associated with the frequency and position of interactions relative to the network core, so that the network consisted of a persistent core of frequent interactions and many peripheral, infrequent interactions. Null model analyses suggest that species abundances play a substantial role in generating these patterns. Our study represents an important step in the study of ecological networks, furthering our mechanistic understanding of the ecological processes driving the temporal persistence of interactions.

Intraspecific trait variation and the leaf economics spectrum across resource gradients and levels of organization.

Abstract: Understanding patterns of functional trait variation across environmental gradients offers an opportunity to increase inference in the mechanistic causes of plant community assembly. The leaf economics spectrum (LES) predicts global tradeoffs in leaf traits and trait-environment relationships, but few studies have examined whether these predictions hold across different levels of organization, particularly within species. Here, we asked (1) whether the main assumptions of the LES (expected trait relationships and shifts in trait values across resource gradients) hold at the intraspecific level, and (2) how within-species trait correlations scale up to interspecific or among-community levels. We worked with leaf traits of saplings of woody species growing across light and soil N and P availability gradients in temperate rainforests of southern Chile. We found that ITV accounted for a large proportion of community-level variation in leaf traits (e.g., LMA and leaf P) and played an important role in driving community-level shifts in leaf traits across environmental gradients. Additionally, intraspecific leaf trait relationships were generally consistent with interspecific and community-level trait relationships and with LES predictions-e.g., a strong negative intraspecific LMA-leaf N correlation-although, most trait relationships varied significantly among species, suggesting idiosyncrasies in the LES at the intraspecific level.

Can mesophotic reefs replenish shallow reefs? Reduced coral reproductive performance casts a doubt.

Abstract: Mesophotic coral ecosystems (i.e., deep coral reefs at 30-120 m depth) appear to be thriving while many shallow reefs in the world are declining. Amid efforts to understand and manage their decline, it was suggested that mesophotic reefs might serve as natural refuges and a possible source of propagules for the shallow reefs. However, our knowledge of how reproductive performance of corals alters with depth is sparse. Here, we present a comprehensive study of the reproductive phenology, fecundity, and abundance of seven reef-building conspecific corals in shallow and mesophotic habitats. Significant differences were found in the synchrony and timing of gametogenesis and spawning between shallow and mesophotic coral populations. Thus, mesophotic populations exhibited delayed or protracted spawning events, which led to spawning of the mesophotic colonies in large proportions at times where the shallow ones had long been depleted of reproductive material. All species investigated demonstrated a substantial reduction in fecundity and/or oocyte sizes at mesophotic depths (40-60 m). Two species (Seriatopora hystrix and Galaxea fascicularis) displayed a reduction in both fecundity and oocyte size at mesophotic depths. Turbinaria reniformis had only reduced fecundity and Acropora squarrosa and Acropora valida only reduced oocyte size. In Montipora verrucosa, reduced fecundity was found during one annual reproductive season while, in the following year, only reduced oocyte size was found. In contrast, reduced oocyte size in mesophotic populations of Acropora squarrosa was consistent along three studied years. One species, Acropora pharaonis, was found to be infertile at mesophotic depths along two studied years. This indicates that reproductive performance decreases with depth; and that although some species are capable of reproducing at mesophotic depths, their contribution to the replenishment of shallow reefs may be inconsequential. Reduced reproductive performance with depth, combined with the possible narrower tolerance to environmental factors, further suggests that mesophotic corals may in fact be more vulnerable than previously conceived. Furthermore, we posit that the observed temporal segregation in reproduction could lead to assortative mating, and this, in turn, may facilitate adaptive divergence across depth.

Using niche breadth theory to explain generalization in mutualisms

Abstract: For a mutualism to remain evolutionarily stable, theory predicts that mutualists should limit their associations to high-quality partners. However, most mutualists either simultaneously or sequentially associate with multiple partners that confer the same type of reward. By viewing mutualisms through the lens of niche breadth evolution, we outline how the environment shapes partner availability and relative quality, and ultimately a focal mutualist's partner breadth. We argue that mutualists that associate with multiple partners may have a selective advantage compared to specialists for many reasons, including sampling, complementarity, and portfolio effects, as well as the possibility that broad partner breadth increases breadth along other niche axes. Furthermore, selection for narrow partner breadth is unlikely to be strong when the environment erodes variation in partner quality, reduces the costs of interacting with low-quality partners, spatially structures partner communities, or decreases the strength of mutualism. Thus, we should not be surprised that most mutualists have broad partner breadth, even if it allows for ineffective partners to persist.

Increasing connectivity between metapopulation ecology and landscape ecology.

Abstract: Metapopulation ecology and landscape ecology aim to understand how spatial structure influences ecological processes, yet these disciplines address the problem using fundamentally different modeling approaches. Metapopulation models describe how the spatial distribution of patches affects colonization and extinction, but often do not account for the heterogeneity in the landscape between patches. Models in landscape ecology use detailed descriptions of landscape structure, but often without considering colonization and extinction dynamics. We present a novel spatially explicit modeling framework for narrowing the divide between these disciplines to advance understanding of the effects of landscape structure on metapopulation dynamics. Unlike previous efforts, this framework allows for statistical inference on landscape resistance to colonization using empirical data. We demonstrate the approach using 11 yr of data on a threatened amphibian in a desert ecosystem. Occupancy data for Lithobates chiricahuensis (Chiricahua leopard frog) were collected on the Buenos Aires National Wildlife Refuge (BANWR), Arizona, USA from 2007 to 2017 following a reintroduction in 2003. Results indicated that colonization dynamics were influenced by both patch characteristics and landscape structure. Landscape resistance increased with increasing elevation and distance to the nearest streambed. Colonization rate was also influenced by patch quality, with semi-permanent and permanent ponds contributing substantially more to the colonization of neighboring ponds relative to intermittent ponds. Ponds that only hold water intermittently also had the highest extinction rate. Our modeling framework can be widely applied to understand metapopulation dynamics in complex landscapes, particularly in systems in which the environment between habitat patches influences the colonization process.

Invoking adaptation to decipher the genetic legacy of past climate change

Abstract: Persistence of natural populations during periods of climate change is likely to depend on migration (range shifts) or adaptation. These responses were traditionally considered discrete processes and conceptually divided into the realms of ecology and evolution. In a milestone paper, Davis and Shaw (2001) Science 292:673 argued that the interplay of adaptation and migration was central to biotic responses to Quaternary climate, but since then there has been no synthesis of efforts made to set up this research program. Here we review some of the salient findings from molecular genetic studies assessing ecological and evolutionary responses to Quaternary climate change. These studies have revolutionized our understanding of population processes associated with past species migration. However, knowledge remains limited about the role of natural selection for local adaptation of populations to Quaternary environmental fluctuations and associated range shifts, and for the footprints this might have left on extant populations. Next-generation sequencing technologies, high-resolution paleoclimate analyses, and advances in population genetic theory offer an unprecedented opportunity to test hypotheses about adaptation through time. Recent population genomics studies have greatly improved our understanding of the role of contemporary adaptation to local environments in shaping spatial patterns of genetic diversity across modern-day landscapes. Advances in this burgeoning field provide important conceptual and methodological bases to decipher the historical role of natural selection and assess adaptation to past environmental variation. We suggest that a process called "temporal conditional neutrality" has taken place: some alleles favored in glacial environments become selectively neutral in modern-day conditions, whereas some alleles that had been neutral during glacial periods become under selection in modern environments. Building on this view, we present a new integrative framework for addressing the interplay of demographic and adaptive evolutionary responses to Quaternary climate dynamics, the research agenda initially envisioned by Davis and Shaw (2001) Science 292:673.

Interactive effects of climate change and biodiversity loss on ecosystem functioning.

Abstract: Climate change and biodiversity loss are expected to simultaneously affect ecosystems, however research on how each driver mediates the effect of the other has been limited in scope. The multiple stressor framework emphasizes non-additive effects, but biodiversity may also buffer the effects of climate change, and climate change may alter which mechanisms underlie biodiversity-function relationships. Here, we performed an experiment using tank bromeliad ecosystems to test the various ways that rainfall changes and litter diversity may jointly determine ecological processes. Litter diversity and rainfall changes interactively affected multiple functions, but how depends on the process measured. High litter diversity buffered the effects of altered rainfall on detritivore communities, evidence of insurance against impacts of climate change. Altered rainfall affected the mechanisms by which litter diversity influenced decomposition, reducing the importance of complementary attributes of species (complementarity effects), and resulting in an increasing dependence on the maintenance of specific species (dominance effects). Finally, altered rainfall conditions prevented litter diversity from fueling methanogenesis, because such changes in rainfall reduced microbial activity by 58%. Together, these results demonstrate that the effects of climate change and biodiversity loss on ecosystems cannot be understood in isolation and interactions between these stressors can be multifaceted.

Disease where you dine: plant species and floral traits associated with pathogen transmission in bumble bees.

Abstract: Hotspots of disease transmission can strongly influence pathogen spread Bee pathogens may be transmitted via shared floral use, but the role of plant species and floral trait variation in shaping transmission dynamics is almost entirely unexplored Given the importance of pathogens for the decline of several bee species, understanding whether and how plant species and floral traits affect transmission could give us important tools for predicting which plant species may be hotspots for disease spread We assessed variation in transmission via susceptibility (probability of infection) and mean intensity (cell count of infected bees) of the trypanosomatid gut pathogen Crithidia bombi to uninfected Bombus impatiens workers foraging on 14 plant species, and assessed the role of floral traits, bee size and foraging behavior on transmission We also conducted a manipulative experiment to determine how the number of open flowers affected transmission on three plant species, Penstemon digitalis, Monarda didyma, and Lythrum salicaria Plant species differed fourfold in the overall mean abundance of Crithidia in foraging bumble bees (mean including infected and uninfected bees) Across plant species, bee susceptibility and mean intensity increased with the number of reproductive structures per inflorescence (buds, flowers and fruits); smaller bees and those that foraged longer were also more susceptible Trait-based models were as good or better than species-based models at predicting susceptibility and mean intensity based on AIC values Surprisingly, floral size and morphology did not significantly predict transmission across species In the manipulative experiment, more open flowers increased mean pathogen abundance fourfold in Monarda, but had no effect in the other two plant species Our results suggest that variation among plant species, through their influence on pathogen transmission, may shape bee disease dynamics Given widespread investment in pollinator-friendly plantings to support pollinators, understanding how plant species affect disease transmission is important for recommending plant species that optimize pollinator health

Bottom-up linkages between primary production, zooplankton, and fish in a shallow, hypereutrophic lake

Abstract: Nutrient supply is a key bottom-up control of phytoplankton primary production in lake ecosystems. Top-down control via grazing pressure by zooplankton also constrains primary production and primary production may simultaneously affect zooplankton. Few studies have addressed these bidirectional interactions. We used convergent cross-mapping (CCM), a numerical test of causal associations, to quantify the presence and direction of the causal relationships among environmental variables (light availability, surface water temperature, NO3 -N, and PO4 -P), phytoplankton community composition, primary production, and the abundances of five functional zooplankton groups (large cladocerans, small cladocerans, rotifers, calanoids, and cyclopoids) in Lake Kasumigaura, a shallow, hypereutrophic lake in Japan. CCM suggested that primary production was causally influenced by NO3 -N and phytoplankton community composition; there was no detectable evidence of a causal effect of zooplankton on primary production. Our results also suggest that rotifers and cyclopoids were forced by primary production, and cyclopoids were further influenced by rotifers. However, our CCM suggested that primary production was weakly influenced by rotifers (i.e., bidirectional interaction). These findings may suggest complex linkages between nutrients, primary production, and rotifers and cyclopoids, a pattern that has not been previously detected or has been neglected. We used linear regression analysis to examine the relationships between the zooplankton community and pond smelt (Hypomesus nipponensis), the most abundant planktivore and the most important commercial fish species in Lake Kasumigaura. The relative abundance of pond smelt was significantly and positively correlated with the abundances of rotifers and cyclopoids, which were causally influenced by primary production. This finding suggests that bottom-up linkages between nutrient, primary production, and zooplankton abundance might be a key mechanism supporting high planktivore abundance in eutrophic lakes. Because increases in primary production and cyanobacteria blooms are likely to occur simultaneously in hypereutrophic lakes, our study highlights the need for ecosystem management to resolve the conflict between good water quality and high fishery production.