Showing papers in "Ecology in 2016"

The ecological impacts of marine debris: unraveling the demonstrated evidence from what is perceived.

Abstract: Anthropogenic debris contaminates marine habitats globally, leading to several perceived ecological impacts. Here, we critically and systematically review the literature regarding impacts of debris from several scientific fields to understand the weight of evidence regarding the ecological impacts of marine debris. We quantified perceived and demonstrated impacts across several levels of biological organization that make up the ecosystem and found 366 perceived threats of debris across all levels. Two hundred and ninety-six of these perceived threats were tested, 83% of which were demonstrated. The majority (82%) of demonstrated impacts were due to plastic, relative to other materials (e.g., metals, glass) and largely (89%) at suborganismal levels (e.g., molecular, cellular, tissue). The remaining impacts, demonstrated at higher levels of organization (i.e., death to individual organisms, changes in assemblages), were largely due to plastic marine debris (> 1 mm; e.g., rope, straws, and fragments). Thus, we show evidence of ecological impacts from marine debris, but conclude that the quantity and quality of research requires improvement to allow the risk of ecological impacts of marine debris to be determined with precision. Still, our systematic review suggests that sufficient evidence exists for decision makers to begin to mitigate problematic plastic debris now, to avoid risk of irreversible harm.

Hydrological and biogeochemical controls on watershed dissolved organic matter transport: pulse-shunt concept

Abstract: Hydrological precipitation and snowmelt events trigger large "pulse" releases of terrestrial dissolved organic matter (DOM) into drainage networks due to an increase in DOM concentration with discharge. Thus, low-frequency large events, which are predicted to increase with climate change, are responsible for a significant percentage of annual terrestrial DOM input to drainage networks. These same events are accompanied by marked and rapid increases in headwater stream velocity; thus they also "shunt" a large proportion of the pulsed DOM to downstream, higher-order rivers and aquatic ecosystems geographically removed from the DOM source of origin. Here we merge these ideas into the "pulse-shunt concept" (PSC) to explain and quantify how infrequent, yet major hydrologic events may drive the timing, flux, geographical dispersion, and regional metabolism of terrestrial DOM. The PSC also helps reconcile long-standing discrepancies in C cycling theory and provides a robust framework for better quantifying its highly dynamic role in the global C cycle. The PSC adds a critical temporal dimension to linear organic matter removal dynamics postulated by the river continuum concept. It also can be represented mathematically through a model that is based on stream scaling approaches suitable for quantifying the important role of streams and rivers in the global C cycle. Initial hypotheses generated by the PSC include: (1) Infrequent large storms and snowmelt events account for a large and underappreciated percentage of the terrestrial DOM flux to drainage networks at annual and decadal time scales and therefore event statistics are equally important to total discharge when determining terrestrial fluxes. (2) Episodic hydrologic events result in DOM bypassing headwater streams and being metabolized in large rivers and exported to coastal systems. We propose that the PSC provides a framework for watershed biogeochemical modeling and predictions and discuss implications to ecological processes.

Hierarchical filters determine community assembly of urban species pools

Abstract: The majority of humanity now lives in cities or towns, with this proportion expected to continue increasing for the foreseeable future. As novel ecosystems, urban areas offer an ideal opportunity to examine multi-scalar processes involved in community assembly as well as the role of human activities in modulating environmental drivers of biodiversity. Although ecologists have made great strides in recent decades at documenting ecological relationships in urban areas, much remains unknown, and we still need to identify the major ecological factors, aside from habitat loss, behind the persistence or extinction of species and guilds of species in cities. Given this paucity of knowledge, there is an immediate need to facilitate collaborative, interdisciplinary research on the patterns and drivers of biodiversity in cities at multiple spatial scales. In this review, we introduce a new conceptual framework for understanding the filtering processes that mold diversity of urban floras and faunas. We hypothesize that the following hierarchical series of filters influence species distributions in cities: (1) regional climatic and biogeographical factors; (2) human facilitation; (3) urban form and development history; (4) socioeconomic and cultural factors; and (5) species interactions. In addition to these filters, life history and functional traits of species are important in determining community assembly and act at multiple spatial scales. Using these filters as a conceptual framework can help frame future research needed to elucidate processes of community assembly in urban areas. Understanding how humans influence community structure and processes will aid in the management, design, and planning of our cities to best support biodiversity.

Estimating local biodiversity change: a critique of papers claiming no net loss of local diversity.

Abstract: Global species extinction rates are orders of magnitude above the background rate documented in the fossil record. However, recent data syntheses have found mixed evidence for patterns of net species loss at local spatial scales. For example, two recent data meta-analyses have found that species richness is decreasing in some locations and is increasing in others. When these trends are combined, these papers argued there has been no net change in species richness, and suggested this pattern is globally representative of biodiversity change at local scales. Here we reanalyze results of these data syntheses and outline why this conclusion is unfounded. First, we show the datasets collated for these syntheses are spatially biased and not representative of the spatial distribution of species richness or the distribution of many primary drivers of biodiversity change. This casts doubt that their results are representative of global patterns. Second, we argue that detecting the trend in local species richness is very difficult with short time series and can lead to biased estimates of change. Reanalyses of the data detected a signal of study duration on biodiversity change, indicating net biodiversity loss is most apparent in studies of longer duration. Third, estimates of species richness change can be biased if species gains during post-disturbance recovery are included without also including species losses that occurred during the disturbance. Net species gains or losses should be assessed with respect to common baselines or reference communities. Ultimately, we need a globally coordinated effort to monitor biodiversity so that we can estimate and attribute human impacts as causes of biodiversity change. A combination of technologies will be needed to produce regularly updated global datasets of local biodiversity change to guide future policy. At this time the conclusion that there is no net change in local species richness is not the consensus state of knowledge.

Unifying error structures in commonly used biotracer mixing models

Abstract: Mixing models are statistical tools that use biotracers to probabilistically estimate the contribution of multiple sources to a mixture. These biotracers may include contaminants, fatty acids, or stable isotopes, the latter of which are widely used in trophic ecology to estimate the mixed diet of consumers. Bayesian implementations of mixing models using stable isotopes (e.g., MixSIR, SIAR) are regularly used by ecologists for this purpose, but basic questions remain about when each is most appropriate. In this study, we describe the structural differences between common mixing model error formulations in terms of their assumptions about the predation process. We then introduce a new parameterization that unifies these mixing model error structures, as well as implicitly estimates the rate at which consumers sample from source populations (i.e., consumption rate). Using simulations and previously published mixing model datasets, we demonstrate that the new error parameterization outperforms existing models and provides an estimate of consumption. Our results suggest that the error structure introduced here will improve future mixing model estimates of animal diet.

Heterogeneity in ecological and evolutionary meta‐analyses: its magnitude and implications

Abstract: Meta-analysis is the gold standard for synthesis in ecology and evolution. Together with estimating overall effect magnitudes, meta-analyses estimate differences between effect sizes via heterogeneity statistics. It is widely hypothesized that heterogeneity will be present in ecological/evolutionary meta-analyses due to the system-specific nature of biological phenomena. Despite driving recommended best practices, the generality of heterogeneity in ecological data has never been systematically reviewed. We reviewed 700 studies, finding 325 that used formal meta-analysis, of which total heterogeneity was reported in fewer than 40%. We used second-order meta-analysis to collate heterogeneity statistics from 86 studies. Our analysis revealed that the median and mean heterogeneity, expressed as I2 , are 84.67% and 91.69%, respectively. These estimates are well above "high" heterogeneity (i.e., 75%), based on widely adopted benchmarks. We encourage reporting heterogeneity in the forms of I2 and the estimated variance components (e.g., τ2 ) as standard practice. These statistics provide vital insights in to the degree to which effect sizes vary, and provide the statistical support for the exploration of predictors of effect-size magnitude. Along with standard meta-regression techniques that fit moderator variables, multi-level models now allow partitioning of heterogeneity among correlated (e.g., phylogenetic) structures that exist within data.

A novel soil manganese mechanism drives plant species loss with increased nitrogen deposition in a temperate steppe

Abstract: Loss of plant diversity with increased anthropogenic nitrogen (N) deposition in grasslands has occurred globally. In most cases, competitive exclusion driven by preemption of light or space is invoked as a key mechanism. Here, we provide evidence from a 9-yr N-addition experiment for an alternative mechanism: differential sensitivity of forbs and grasses to increased soil manganese (Mn) levels. In Inner Mongolia steppes, increasing the N supply shifted plant community composition from grass-forb codominance (primarily Stipa krylovii and Artemisia frigida, respectively) to exclusive dominance by grass, with associated declines in overall species richness. Reduced abundance of forbs was linked to soil acidification that increased mobilization of soil Mn, with a 10-fold greater accumulation of Mn in forbs than in grasses. The enhanced accumulation of Mn in forbs was correlated with reduced photosynthetic rates and growth, and is consistent with the loss of forb species. Differential accumulation of Mn between forbs and grasses can be linked to fundamental differences between dicots and monocots in the biochemical pathways regulating metal transport. These findings provide a mechanistic explanation for N-induced species loss in temperate grasslands by linking metal mobilization in soil to differential metal acquisition and impacts on key functional groups in these ecosystems.

Structural complexity and component type increase intertidal biodiversity independently of area

Abstract: Complexity is well accepted as one of the primary drivers of biodiversity, however, empirical support for such positive associations is often confounded with surface area and undermined by other potential explanatory factors, especially the type of structural component (e.g., pits, crevices, overhangs, etc.). In the present study, sample units (artificial substrates) of equal surface area (± 0.2%) were used to simultaneously examine the independent effects of complexity and different structural component types on species richness (S), abundance (N), and community composition. We created simple and complex concrete substrates of four different geometric designs using novel software. The substrates (n = 8) were mounted onto granite seawalls (at two tidal heights) on two islands south of Singapore Island. After 13 months of colonization, all 384 tiles were collected and their assemblages compared. A total of 53 744 individuals of 70 species/morphospecies were collected and identified. Our results show that greater complexity can support greater species richness and different communities that are independent of surface area. Furthermore, the type of structure (e.g., "pits," "grooves," "towers") can have an effect on richness and community composition that is independent of complexity.

Resource waves: phenological diversity enhances foraging opportunities for mobile consumers

Abstract: Time can be a limiting constraint for consumers, particularly when resource phenology mediates foraging opportunity. Though a large body of research has explored how resource phenology influences trophic interactions, this work has focused on the topics of trophic mismatch or predator swamping, which typically occur over short periods, at small spatial extents or coarse resolutions. In contrast many consumers integrate across landscape heterogeneity in resource phenology, moving to track ephemeral food sources that propagate across space as resource waves. Here we provide a conceptual framework to advance the study of phenological diversity and resource waves. We define resource waves, review evidence of their importance in recent case studies, and demonstrate their broader ecological significance with a simulation model. We found that consumers ranging from fig wasps (Chalcidoidea) to grizzly bears (Ursus arctos) exploit resource waves, integrating across phenological diversity to make resource aggregates available for much longer than their component parts. In model simulations, phenological diversity was often more important to consumer energy gain than resource abundance per se. Current ecosystem-based management assumes that species abundance mediates the strength of trophic interactions. Our results challenge this assumption and highlight new opportunities for conservation and management. Resource waves are an emergent property of consumer-resource interactions and are broadly significant in ecology and conservation.

Plant functional traits of dominant native and invasive species in Mediterranean-climate ecosystems

Abstract: The idea that dominant invasive plant species outperform neighboring native species through higher rates of carbon assimilation and growth is supported by several analyses of global data sets. However, theory suggests that native and invasive species occurring in low-resource environments will be functionally similar, as environmental factors restrict the range of observed physiological and morphological trait values. We measured resource-use traits in native and invasive plant species across eight diverse vegetation communities distributed throughout the five mediterranean-climate regions, which are drought prone and increasingly threatened by human activities, including the introduction of exotic species. Traits differed strongly across the five regions. In regions with functional differences between native and invasive species groups, invasive species displayed traits consistent with high resource acquisition; however, these patterns were largely attributable to differences in life form. We found that species invading mediterranean-climate regions were more likely to be annual than perennial: three of the five regions were dominated by native woody species and invasive annuals. These results suggest that trait differences between native and invasive species are context dependent and will vary across vegetation communities. Native and invasive species within annual and perennial groups had similar patterns of carbon assimilation and resource use, which contradicts the widespread idea that invasive species optimize resource acquisition rather than resource conservation.

Generalist birds promote tropical forest regeneration and increase plant diversity via rare‐biased seed dispersal

Abstract: Regenerated forests now compose over half of the world's tropical forest cover and are increasingly important as providers of ecosystem services, freshwater, and biodiversity conservation. Much of the value and functionality of regenerating forests depends on the plant diversity they contain. Tropical forest diversity is strongly shaped by mutualistic interactions between plants and fruit-eating animals (frugivores) that disperse seeds. Here we show how seed dispersal by birds can influence the speed and diversity of early successional forests in Puerto Rico. For two years, we monitored the monthly fruit production of bird-dispersed plants on a fragmented landscape, and measured seed dispersal activity of birds and plant establishment in experimental plots located in deforested areas. Two predominantly omnivorous bird species, the Northern Mockingbird (Mimus polyglottos) and the Gray Kingbird (Tyrannus dominicensis), proved critical for speeding up the establishment of woody plants and increasing the species richness and diversity of the seed rain in deforested areas. Seed dispersal by these generalists increased the odds for rare plant species to disperse and establish in experimental forest-regeneration plots. Results indicate that birds that mix fruit and insects in their diets and actively forage across open and forested habitats can play keystone roles in the regeneration of mutualistic plant-animal communities. Furthermore, our analyses reveal that rare-biased (antiapostatic) frugivory and seed dispersal is the mechanism responsible for increasing plant diversity in the early-regenerating community.

The importance of biodiversity and dominance for multiple ecosystem functions in a human-modified tropical landscape.

Abstract: Many studies suggest that biodiversity may be particularly important for ecosystem multifunctionality, because different species with different traits can contribute to different functions. Support, however, comes mostly from experimental studies conducted at small spatial scales in low-diversity systems. Here, we test whether different species contribute to different ecosystem functions that are important for carbon cycling in a high-diversity human-modified tropical forest landscape in Southern Mexico. We quantified aboveground standing biomass, primary productivity, litter production, and wood decomposition at the landscape level, and evaluated the extent to which tree species contribute to these ecosystem functions. We used simulations to tease apart the effects of species richness, species dominance and species functional traits on ecosystem functions. We found that dominance was more important than species traits in determining a species' contribution to ecosystem functions. As a consequence of the high dominance in human-modified landscapes, the same small subset of species mattered across different functions. In human-modified landscapes in the tropics, biodiversity may play a limited role for ecosystem multifunctionality due to the potentially large effect of species dominance on biogeochemical functions. However, given the spatial and temporal turnover in species dominance, biodiversity may be critically important for the maintenance and resilience of ecosystem functions.

Predicting nutrient excretion of aquatic animals with metabolic ecology and ecological stoichiometry: a global synthesis

Abstract: The metabolic theory of ecology (MTE) and ecological stoichiometry (ES) are both prominent frameworks for understanding energy and nutrient budgets of organisms. We tested their separate and joint power to predict nitrogen (N) and phosphorus (P) excretion rates of ectothermic aquatic invertebrate and vertebrate animals (10,534 observations worldwide). MTE variables (body size, temperature) performed better than ES variables (trophic guild, vertebrate classification, body N:P) in predicting excretion rates, but the best models included variables from both frameworks. Size scaling coefficients were significantly lower than predicted by MTE (<0.75), were lower for P than N, and varied greatly among species. Contrary to expectations under ES, vertebrates excreted both N and P at higher rates than invertebrates despite having more nutrient-rich bodies, and primary consumers excreted as much nutrients as carnivores despite having nutrient-poor diets. Accounting for body N:P hardly improved upon predictions from treating vertebrate classification categorically. We conclude that basic data on body size, water temperature, trophic guild, and vertebrate classification are sufficient to make general estimates of nutrient excretion rates for any animal taxon or aquatic ecosystem. Nonetheless, dramatic interspecific variation in size-scaling coefficients and counter-intuitive patterns with respect to diet and body composition underscore the need for field data on consumption and egestion rates. Together, MTE and ES provide a powerful conceptual basis for interpreting and predicting nutrient recycling rates of aquatic animals worldwide.

Why do trees die? Characterizing the drivers of background tree mortality.

Abstract: The drivers of background tree mortality rates—the typical low rates of tree mortality found in forests in the absence of acute stresses like drought—are central to our understanding of forest dynamics, the effects of ongoing environmental changes on forests, and the causes and consequences of geographical gradients in the nature and strength of biotic interactions. To shed light on factors contributing to background tree mortality, we analyzed detailed pathological data from 200,668 tree-years of observation and 3,729 individual tree deaths, recorded over a 13-yr period in a network of old-growth forest plots in California's Sierra Nevada mountain range. We found that: (1) Biotic mortality factors (mostly insects and pathogens) dominated (58%), particularly in larger trees (86%). Bark beetles were the most prevalent (40%), even though there were no outbreaks during the study period; in contrast, the contribution of defoliators was negligible. (2) Relative occurrences of broad classes of mortality factors (biotic, 58%; suppression, 51%; and mechanical, 25%) are similar among tree taxa, but may vary with tree size and growth rate. (3) We found little evidence of distinct groups of mortality factors that predictably occur together on trees. Our results have at least three sets of implications. First, rather than being driven by abiotic factors such as lightning or windstorms, the “ambient” or “random” background mortality that many forest models presume to be independent of tree growth rate is instead dominated by biotic agents of tree mortality, with potentially critical implications for forecasting future mortality. Mechanistic models of background mortality, even for healthy, rapidly growing trees, must therefore include the insects and pathogens that kill trees. Second, the biotic agents of tree mortality, instead of occurring in a few predictable combinations, may generally act opportunistically and with a relatively large degree of independence from one another. Finally, beyond the current emphasis on folivory and leaf defenses, studies of broad-scale gradients in the nature and strength of biotic interactions should also include biotic attacks on, and defenses of, tree stems and roots.

The contribution of germination functional traits to population dynamics of a desert plant community.

Abstract: Early life-cycle events play critical roles in determining the population and community dynamics of plants. The ecology of seeds and their germination patterns can determine range limits, adaptation to environmental variation, species diversity, and community responses to climate change. Understanding the adaptive consequences and environmental filtering of such functional traits will allow us to explain and predict ecological dynamics. Here we quantify key functional aspects of germination physiology and relate them to an existing functional ecology framework to explain long-term population dynamics for 13 species of desert annuals near Tucson, Arizona, USA. Our goal was to assess the extent to which germination functional biology contributes to long-term population processes in nature. Some of the species differences in base, optimum, and maximum temperatures for germination, thermal times to germination, and base water potentials for germination were strongly related to 20-yr mean germination fractions, 25-yr average germination dates, seed size, and long-term demographic variation. Comparisons of germination fraction, survival, and fecundity vs. yearly changes in population size found significant roles for all three factors, although in varying proportions for different species. Relationships between species' germination physiologies and relative germination fractions varied across years, with fast-germinating species being favored in years with warm temperatures during rainfall events in the germination season. Species with low germination fractions and high demographic variance have low integrated water-use efficiency, higher vegetative growth rates, and smaller, slower-germinating seeds. We have identified and quantified a number of functional traits associated with germination biology that play critical roles in ecological population dynamics.

Nitrogen addition affects chemical compositions of plant tissues, litter and soil organic matter.

Abstract: Increasing nitrogen (N) deposition or fertilization has been found to significantly affect carbon (C) cycling. However, a comprehensive understanding of how different C chemical components of plant, litter, and soil would respond to external N addition is still lacking. We compiled data of 1,160 observations from 52 individual studies and conducted a meta-analysis of N addition effects on 18 variables related to C chemical compositions in terrestrial ecosystems. Results showed that plant lignin (+7.13%), plant protein (+25.94%), and soil lignin (+7.30%) were significantly increased by N addition, and plant hemicellulose (-4.39%) was significantly decreased, whereas plant fiber, plant cellulose, plant non-structural carbohydrate (NSC), litter lignin, and litter cellulose were not significantly changed. The effects of N addition on C chemical composition varied among different ecosystems/plant types and different forms of N addition. Increasing treatment duration did not significantly change the effects of N addition on the chemical composition of plant, litter, and soil C. With increasing N addition rate, the effect of N addition on plant lignin, plant fiber, plant cellulose, and plant protein increased, while the effect of N addition on plant hemicellulose, plant NSC, and litter cellulose became more negative. Our meta-analysis provided a systematic evaluation of the responses of different C chemical components to N addition in the plant-litter-soil continuum. Results suggest that the change of plant and soil C chemical composition under N addition may be beneficial for ecosystem C sequestration and could affect ecosystem structure and function in the future.

Phylogenetic impoverishment of plant communities following chronic human disturbances in the Brazilian Caatinga.

Abstract: Chronic disturbances, such as selective logging, firewood extraction and extensive grazing, may lead to the taxonomic and phylogenetic impoverishment of remaining old-growth forest communities worldwide; however, the empirical evidence on this topic is limited. We tested this hypothesis in the Caatinga vegetation--a seasonally dry tropical forest restricted to northeast Brazil. We sampled 11,653 individuals (adults, saplings, and seedlings) from 51 species in 29 plots distributed along a gradient of chronic disturbance. The gradient was assessed using a chronic disturbance index (CDI) based on five recognized indicators of chronic disturbances: proximity to urban center, houses and roads and the density of both people and livestock. We used linear models to test if mean effective number of lineages, mean phylogenetic distance and phylogenetic dispersion decreased with CDI and if such relationships differed among ontogenetic stages. As expected, the mean effective number of lineages and the mean phylogenetic distance were negatively related to CDI, and such diversity losses occurred irrespective of ontogeny. Yet the increase in phylogenetic clustering in more disturbed plots was only evident in seedlings and saplings, mostly because clades with more descendent taxa than expected by chance (e.g., Euphorbiaceae) thrived in more disturbed plots. This novel study indicates that chronic human disturbances are promoting the phylogenetic impoverishment of the irreplaceable woody flora of the Brazilian Caatinga forest. The highest impoverishment was observed in seedlings and saplings, indicating that if current chronic disturbances remain, they will result in increasingly poorer phylogenetically forests. This loss of evolutionary history will potentially limit the capacity of this ecosystem to respond to human disturbances (i.e., lower ecological resilience) and particularly their ability to adapt to rapid climatic changes in the region.

Climate and edaphic controllers influence rhizosphere community assembly for a wild annual grass.

Abstract: The interface between roots and soil, known as the rhizosphere, is a dynamic habitat in the soil ecosystem. Unraveling the factors that control rhizosphere community assembly is a key starting point for understanding the diversity of plant-microbial interactions that occur in soil. The goals of this study were to determine how environmental factors shape rhizosphere microbial communities, such as local soil characteristics and the regional climate, and to determine the relative influence of the rhizosphere on microbial community assembly compared to the pressures imposed by the local and regional environment. We identified the bacteria present in the soil immediately adjacent to the roots of wild oat (A vena spp.) in three California grasslands using deep Illumina 16S sequencing. Rhizosphere communities were more similar to each other than to the surrounding soil communities from which they were derived, despite the fact that the grasslands studied were separated by hundreds of kilometers. The rhizosphere was the dominant factor structuring bacterial community composition (38% variance explained), and was comparable in magnitude to the combined local and regional effects (22% and 21%, respectively). Rhizosphere communities were most influenced by factors related to the regional climate (soil moisture and temperature), while background soil communities were more influenced by soil characteristics (pH, CEC, exchangeable cations, clay content). The Avena core microbiome was strongly phylogenetically clustered according to the metrics NRI and NTI, which indicates that selective processes likely shaped these communities. Furthermore, 17% of these taxa were not detectable in the background soil, even with a robust sequencing depth of approximately 70,000 sequences per sample. These results support the hypothesis that roots select less abundant or possibly rare populations in the soil microbial community, which appear to be lineages of bacteria that have made a physiological tradeoff for rhizosphere competence at the expense of their competitiveness in non-rhizosphere soil.

Model selection and assessment for multi-species occupancy models

Abstract: While multi-species occupancy models (MSOMs) are emerging as a popular method for analyzing biodiversity data, formal checking and validation approaches for this class of models have lagged behind. Concurrent with the rise in application of MSOMs among ecologists, a quiet regime shift is occurring in Bayesian statistics where predictive model comparison approaches are experiencing a resurgence. Unlike single-species occupancy models that use integrated likelihoods, MSOMs are usually couched in a Bayesian framework and contain multiple levels. Standard model checking and selection methods are often unreliable in this setting and there is only limited guidance in the ecological literature for this class of models. We examined several different contemporary Bayesian hierarchical approaches for checking and validating MSOMs and applied these methods to a freshwater aquatic study system in Colorado, USA, to better understand the diversity and distributions of plains fishes. Our findings indicated distinct differences among model selection approaches, with cross-validation techniques performing the best in terms of prediction.

Kodiak Brown Bears Surf the Salmon Red Wave: Direct Evidence from GPS Collared Individuals

Abstract: A key constraint faced by consumers is achieving a positive energy balance in the face of temporal variation in foraging opportunities. Recent work has shown that spatial heterogeneity in resource phenology can buffer mobile consumers from this constraint by allowing them to track changes in resource availability across space. For example, salmon populations spawn asynchronously across watersheds, causing high-quality foraging opportunities to propagate across the landscape, prolonging the availability of salmon at the regional scale. However, we know little about how individual consumers integrate across phenological variation or the benefits they receive by doing so. Here, we present direct evidence that individual brown bears track spatial variation in salmon phenology. Data from 40 GPS collared brown bears show that bears visited multiple spawning sites in synchrony with the order of spawning phenology. The number of sites used was correlated with the number of days a bear exploited salmon, suggesting the phenological variation in the study area influenced bear access to salmon, a resource which strongly influences bear fitness. Fisheries managers attempting to maximize harvest while maintaining ecosystem function should strive to protect the population diversity that underlies the phenological variation used by wildlife consumers.

Nectar microbes can reduce secondary metabolites in nectar and alter effects on nectar consumption by pollinators.

Abstract: Secondary metabolites that are present in floral nectar have been hypothesized to enhance specificity in plant-pollinator mutualism by reducing larceny by non-pollinators, including microorganisms that colonize nectar. However, few studies have tested this hypothesis. Using synthetic nectar, we conducted laboratory and field experiments to examine the effects of five chemical compounds found in nectar on the growth and metabolism of nectar-colonizing yeasts and bacteria, and the interactive effects of these compounds and nectar microbes on the consumption of nectar by pollinators. In most cases, focal compounds inhibited microbial growth, but the extent of these effects depended on compound identity, concentration, and microbial species. Moreover, most compounds did not substantially decrease sugar metabolism by microbes, and microbes reduced the concentration of some compounds in nectar. Using artificial flowers in the field, we also found that the common nectar yeast Metschnikowia reukaufii altered nectar consumption by small floral visitors, but only in nectar containing catalpol. This effect was likely mediated by a mechanism independent of catalpol metabolism. Despite strong compound-specific effects on microbial growth, our results suggest that the secondary metabolites tested here are unlikely to be an effective general defense mechanism for preserving nectar sugars for pollinators. Instead, our results indicate that microbial colonization of nectar could reduce the concentration of secondary compounds in nectar and, in some cases, reduce deterrence to pollinators.

Drought, pollen and nectar availability, and pollination success

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Land use intensification in the humid tropics increased both alpha and beta diversity of soil bacteria

Abstract: Anthropogenic pressures on tropical forests are rapidly intensifying, but our understanding of their implications for biological diversity is still very limited, especially with regard to soil biota, and in particular soil bacterial communities. Here we evaluated bacterial community composition and diversity across a gradient of land use intensity in the eastern Amazon from undisturbed primary forest, through primary forests varyingly disturbed by fire, regenerating secondary forest, pasture, and mechanized agriculture. Soil bacteria were assessed by paired-end Illumina sequencing of 16S rRNA gene fragments (V4 region). The resulting sequences were clustered into operational taxonomic units (OTU) at a 97% similarity threshold. Land use intensification increased the observed bacterial diversity (both OTU richness and community heterogeneity across space) and this effect was strongly associated with changes in soil pH. Moreover, land use intensification and subsequent changes in soil fertility, especially pH, altered the bacterial community composition, with pastures and areas of mechanized agriculture displaying the most contrasting communities in relation to undisturbed primary forest. Together, these results indicate that tropical forest conversion impacts soil bacteria not through loss of diversity, as previously thought, but mainly by imposing marked shifts on bacterial community composition, with unknown yet potentially important implications for ecological functions and services performed by these communities.

Burn me twice, shame on who? Interactions between successive forest fires across a temperate mountain region

Abstract: Increasing rates of natural disturbances under a warming climate raise important questions about how multiple disturbances interact. Escalating wildfire activity in recent decades has resulted in some forests re-burning in short succession, but how the severity of one wildfire affects that of a subsequent wildfire is not fully understood. We used a field-validated, satellite-derived, burn-severity atlas to assess interactions between successive wildfires across the US Northern Rocky Mountains a 300,000-km2 region dominated by fire-prone forests. In areas that experienced two wildfires between 1984 and 2010, we asked: (1) How do overall frequency distributions of burn-severity classes compare between first and second fires? (2) In a given location, how does burn severity of the second fire relate to that of the first? (3) Do interactions between successive fires vary by forest zone or the interval between fires? (4) What factors increase the probability of burning twice as stand-replacing fire? Within the study area, 138,061 ha burned twice between 1984 and 2010. Overall, frequency distributions of burn severity classes (low, moderate, high; quantified using relativized remote sensing indices) were similar between the first and second fires; however burn severity was 5–13% lower in second fires on average. Negative interactions between fires were most pronounced in lower-elevation forests and woodlands, when fire intervals were <10 yr, and when burn severity was low in the first fire. When the first fire burned as high severity and fire intervals exceeded 10–12 yr, burn-severity interactions switched from negative to positive, with high-severity fire begetting subsequent high-severity fire. Locations most likely to experience successive stand-replacing fires were high-elevation forests, which are adapted to high-severity fire, and areas conducive to abundant post-fire tree regeneration. Broadly similar severities among short-interval “re-burns” and other wildfires indicate that positive severity feedbacks, an oft-posited agent of ecosystem decline or state shift, are not an inevitable outcome of re-burning. Nonetheless, context-dependent shifts in both the magnitude and direction of wildfire interactions (associated with forest zone, initial burn-severity, and disturbance interval) illustrate complexities in disturbance interactions and can inform management and predictions of future system dynamics.

Inferring species interactions from co-occurrence data with Markov networks.

Abstract: Inferring species interactions from co-occurrence data is one of the most controversial tasks in community ecology. One difficulty is that a single pairwise Interaction can ripple through an ecological network and produce surprising indirect consequences. For example, the negative correlation between two competing species can be reversed in the presence of a third species that outcompetes both of them. Here, I apply models from statistical physics, called Markov networks or Markov random fields, that can predict the direct and indirect consequences of any possible species interaction matrix. Interactions in these models can be estimated from observed co-occurrence rates via maximum likelihood, controlling for indirect effects. Using simulated landscapes with known interactions, I evaluated Markov networks and six existing approaches. Markov networks consistently outperformed the other methods, correctly isolating direct interactions between species pairs even when indirect interactions or abiotic factors largely overpowered them. Two computationally efficient approximations, which controlled for indirect effects with partial correlations or generalized linear models, also performed well. Null models showed no evidence of being able to control for indirect effects, and reliably yielded incorrect inferences when such effects were present. This article is protected by copyright. All rights reserved.

Balls, cups, and quasi-potentials: quantifying stability in stochastic systems.

Abstract: When a system has more than one stable state, how can the stability of these states be compared? This deceptively simple question has important consequences for ecosystems, because systems with alternative stable states can undergo dramatic regime shifts. The probability, frequency, duration, and dynamics of these shifts will all depend on the relative stability of the stable states. Unfortunately, the concept of "stability" in ecology has suffered from substantial confusion and this is particularly problematic for systems where stochastic perturbations can cause shifts between coexisting alternative stable states. A useful way to visualize stable states in stochastic systems is with a ball-in-cup-diagram, in which the state of the system is represented as the position of a ball rolling on a surface, and the random perturbations can push the ball from one basin of attraction to another. The surface is determined by a potential function, which provides a natural stability metric. Systems amenable to this representation, called gradient systems, are quite rare, however. As a result, the potential function is not widely used and other approaches based on linear stability analysis have become standard. Linear stability analysis is designed for local analysis of deterministic systems and, as we show, can produce a highly misleading picture of how the system will behave under continual, stochastic perturbations. In this paper, we show how the potential function can be generalized so that it can be applied broadly, employing a concept from stochastic analysis called the quasi-potential. Using three classic ecological models, we demonstrate that the quasi-potential provides a useful way to quantify stability in stochastic systems. We show that the quasi-potential framework helps clarify long-standing confusion about stability in stochastic ecological systems, and we argue that ecologists should adopt it as a practical tool for analyzing these systems.

Global patterns and determinants of forest canopy height.

Abstract: Forest canopy height is an important indicator of forest biomass, species diversity, and other ecosystem functions; however, the climatic determinants that underlie its global patterns have not been fully explored. Using satellite LiDAR-derived forest canopy heights and field measurements of the world's giant trees, combined with climate indices, we evaluated the global patterns and determinants of forest canopy height. The mean canopy height was highest in tropical regions, but tall forests (>50 m) occur at various latitudes. Water availability, quantified by the difference between annual precipitation and annual potential evapotranspiration (P−PET), was the best predictor of global forest canopy height, which supports the hydraulic limitation hypothesis. However, in striking contrast with previous studies, the canopy height exhibited a hump-shaped curve along a gradient of P−PET: it initially increased, then peaked at approximately 680 mm of P−PET, and finally declined, which suggests that excessive water supply negatively affects the canopy height. This trend held true across continents and forest types, and it was also validated using forest inventory data from China and the United States. Our findings provide new insights into the climatic controls of the world's giant trees and have important implications for forest management and improvement of forest growth models.

Increased stem density and competition may diminish the positive effects of warming at alpine treeline

Abstract: The most widespread response to global warming among alpine treeline ecotones is not an upward shift, but an increase in tree density. However, the impact of increasing density on interactions among trees at treeline is not well understood. Here, we test if treeline densification induced by climatic warming leads to increasing intraspecific competition. We mapped and measured the size and age of Smith fir trees growing in two treelines located in the southeastern Tibetan Plateau. We used spatial point-pattern and codispersion analyses to describe the spatial association and covariation among seedlings, juveniles, and adults grouped in 30-yr age classes from the 1860s to the present. Effects of competition on tree height and regeneration were inferred from bivariate mark-correlations. Since the 1950s, a rapid densification occurred at both sites in response to climatic warming. Competition between adults and juveniles or seedlings at small scales intensified as density increased. Encroachment negatively affected height growth and further reduced recruitment around mature trees. We infer that tree recruitment at the studied treelines was more cold-limited prior to 1950 and shifted to a less temperature-constrained regime in response to climatic warming. Therefore, the ongoing densification and encroachment of alpine treelines could alter the way climate drives their transitions toward subalpine forests.

Underappreciated problems of low replication in ecological field studies.

Abstract: The cost and difficulty of manipulative field studies makes low statistical power a pervasive issue throughout most ecological subdisciplines. Ecologists are already aware that small sample sizes increase the probability of committing Type II errors. In this article, we address a relatively unknown problem with low power: underpowered studies must overestimate small effect sizes in order to achieve statistical significance. First, we describe how low replication coupled with weak effect sizes leads to Type M errors, or exaggerated effect sizes. We then conduct a meta-analysis to determine the average statistical power and Type M error rate for manipulative field experiments that address important questions related to global change; global warming, biodiversity loss, and drought. Finally, we provide recommendations for avoiding Type M errors and constraining estimates of effect size from underpowered studies.

Can trait patterns along gradients predict plant community responses to climate change

Abstract: Plant functional traits vary consistently along climate gradients and are therefore potential predictors of plant community response to climate change. We test this space-for-time assumption by combining a spatial gradient study with whole-community turf transplantation along temperature and precipitation gradients in a network of 12 grassland sites in Southern Norway. Using data on eight traits for 169 species and annual vegetation censuses of 235 turfs over 5 yr, we quantify trait-based responses to climate change by comparing observed community dynamics in transplanted turfs to field-parameterized null model simulations. Three traits related to species architecture (maximum height, number of dormant meristems, and ramet-ramet connection persistence) varied consistently along spatial temperature gradients and also correlated to changes in species abundances in turfs transplanted to warmer climates. Two traits associated with resource acquisition strategy (SLA, leaf area) increased along spatial temperature gradients but did not correlate to changes in species abundances following warming. No traits correlated consistently with precipitation. Our study supports the hypothesis that spatial associations between plant traits and broad-scale climate variables can be predictive of community response to climate change, but it also suggests that not all traits with clear patterns along climate gradients will necessarily influence community response to an equal degree.