Climate change is poised to alter mountain stream ecosystem processes via organismal phenological shifts.

Climate change is affecting the phenology of organisms and ecosystem processes across a wide range of environments. However, the links between organismal and ecosystem process change in complex communities remain uncertain. In snow-dominated watersheds, snowmelt in the spring and early summer, followed by a long low-flow period, characterizes the natural flow regime of streams and rivers. Here, we examined how earlier snowmelt will alter the phenology of mountain stream organisms and ecosystem processes via an outdoor mesocosm experiment in stream channels in the Eastern Sierra Nevada, California. The low-flow treatment, simulating a 3- to 6-wk earlier return to summer baseflow conditions projected under climate change scenarios in the region, increased water temperature and reduced biofilm production to respiration ratios by 32%. Additionally, most of the invertebrate species explaining community change (56% and 67% of the benthic and emergent taxa, respectively), changed in phenology as a consequence of the low-flow treatment. Further, emergent flux pulses of the dominant insect group (Chironomidae) almost doubled in magnitude, benefitting a generalist riparian predator. Changes in both invertebrate community structure (composition) and functioning (production) were mostly fine-scale, and response diversity at the community level stabilized seasonally aggregated responses. Our study illustrates how climate change in vulnerable mountain streams at the rain-to-snow transition is poised to alter the dynamics of stream food webs via fine-scale changes in phenology-leading to novel predator-prey "matches" or "mismatches" even when community structure and ecosystem processes appear stable at the annual scale.

Exploring potential relationships between acoustic indices and ecosystem functions: a test on insect herbivory.

Biodiversity loss is a global concern. Current technological advances allow the development of novel tools that can monitor biodiversity remotely with minimal disturbance. One example is passive acoustic monitoring (PAM), which involves recording the soundscape of an area using autonomous recording units, and processing these data using acoustic indices, for example, to estimate the diversity of various vocal animal groups. We explored the hypothesis that data obtained through PAM could also be used to study ecosystem functions. Specifically, we investigated the potential relationship between seven commonly used acoustic indices and insect leaf herbivory, measured as total leaf damage and as the damage from three major insect feeding guilds. Herbivory was quantified on seedlings in 13 plots in four subtropical forests in south China, and acoustic data, representing insect acoustic complexity, were obtained by recording the evening soundscapes in those same locations. Herbivory levels correlated positively with the acoustic entropy index, commonly reported as one of the best-performing indices, whose high values indicate higher acoustic complexity, likely due to greater insect diversity. Relationships for specific feeding guilds were moderately stronger for chewers, indicating that the acoustic indices capture some insect groups more than others (e.g., chewers include soniferous taxa such as crickets, whereas miners are mostly silent). Our findings suggest that the use of PAM to monitor ecosystem functions deserves to be explored further, as this is a research field with unexplored potential. Well-designed targeted studies could help us better understand how to best use novel technologies to monitor ecosystem functions.

The significance of partial migration for food web and ecosystem dynamics.

Migration is ubiquitous and can strongly shape food webs and ecosystems. Less familiar, however, is that the majority of life cycle, seasonal and diel migrations in nature are partial migrations: only a fraction of the population migrates while the other individuals remain in their resident ecosystem. Here, we demonstrate different impacts of partial migration rendering it fundamental to our understanding of the significance of migration for food web and ecosystem dynamics. First, partial migration affects the spatiotemporal distribution of individuals and the food web and ecosystem-level processes they drive differently than expected under full migration. Second, whether an individual migrates or not is regularly correlated with morphological, physiological, and/or behavioural traits that shape its food-web and ecosystem-level impacts. Third, food web and ecosystem dynamics can drive the fraction of the population migrating, enabling the potential for feedbacks between the causes and consequences of migration within and across ecosystems. These impacts, individually and in combination, can yield unintuitive effects of migration and drive the dynamics, diversity and functions of ecosystems. By presenting the first full integration of partial migration and trophic (meta-)community and (meta-)ecosystem ecology, we provide a roadmap for studying how migration affects and is affected by ecosystem dynamics in a changing world.

ForestClim-Bioclimatic variables for microclimate temperatures of European forests.

Microclimate research gained renewed interest over the last decade and its importance for many ecological processes is increasingly being recognized. Consequently, the call for high-resolution microclimatic temperature grids across broad spatial extents is becoming more pressing to improve ecological models. Here, we provide a new set of open-access bioclimatic variables for microclimate temperatures of European forests at 25 × 25 m2 resolution.

Crop-pollinating Diptera have diverse diets and habitat needs in both larval and adult stages.

Insects are important pollinators of global food crops and wild plants. The adult and larval diet and habitat needs are well known for many bee taxa, but poorly understood for other pollinating taxa. Non-bee pollinators often feed on different substrates in their larval and adult life stages, and this diet and habitat diversity has important implications for their conservation and management. We reviewed the global literature on crop pollinating Diptera (the true flies) to identify both larval and adult fly diet and habitat needs. We then assembled the published larval and adult diets and habitat needs of beneficial fly pollinators found globally into a freely accessible database. Of the 405 fly species known to visit global food crops, we found relevant published evidence regarding larval and adult diet and habitat information for 254 species, which inhabited all eight global biogeographic regions. We found the larvae of these species lived in 35 different natural habitats and belong to 10 different feeding guilds. Additionally, differences between adult Diptera sexes also impacted diet needs; females from 14 species across five families fed on protein sources other than pollen to start the reproductive process of oogenesis (egg development) while males of the same species fed exclusively on pollen and nectar. While all adult species fed at least partially on floral nectar and/or pollen, only five species were recorded feeding on pollen and no fly larvae fed on nectar. Of the 242 species of larvae with established diet information, 33% were predators (n = 79) and 30% were detritivores (n = 73). Detritivores were the most generalist taxa and utilized 17 different habitats and 12 different feeding substrates. Of all fly taxa, only 2% belonged to the same feeding guild in both active life stages. Our results show that many floral management schemes may be insufficient to support pollinating Diptera. Pollinator conservation strategies in agroecosystems should consider other non-floral resources, such as wet organic materials and dung, as habitats for beneficial fly larvae.

Long-term exclusion of invasive ungulates alters tree recruitment and functional traits but not total forest carbon.

Forests are major carbon (C) sinks, but their ability to sequester C and thus mitigate climate change, varies with the environment, disturbance regime, and biotic interactions. Herbivory by invasive, nonnative ungulates can have profound ecosystem effects, yet its consequences for forest C stocks remain poorly understood. We determined the impact of invasive ungulates on C pools, both above- and belowground (to 30 cm), and on forest structure and diversity using 26 paired long-term (>20 years) ungulate exclosures and adjacent unfenced control plots located in native temperate rainforests across New Zealand, spanning 36-41° S. Total ecosystem C was similar between ungulate exclosure (299.93 ± 25.94 Mg C ha-1 ) and unfenced control (324.60 ± 38.39 Mg C ha-1 ) plots. Most (60%) variation in total ecosystem C was explained by the biomass of the largest tree (mean diameter at breast height [dbh]: 88 cm) within each plot. Ungulate exclusion increased the abundance and diversity of saplings and small trees (dbh ≥2.5, <10 cm) compared with unfenced controls, but these accounted for ~5% of total ecosystem C, demonstrating that a few, large trees dominate the total forest ecosystem C but are unaffected by invasive ungulates at a timescale of 20-50 years. However, changes in understory C pools, species composition, and functional diversity did occur following long-term ungulate exclusion. Our findings suggest that, although the removal of invasive herbivores may not affect total forest C at the decadal scale, major shifts in the diversity and composition of regenerating species will have longer term consequences for ecosystem processes and forest C.

A trait-based framework for dung beetle functional ecology.

Traits are key for understanding the environmental responses and ecological roles of organisms. Trait approaches to functional ecology are well established for plants, whereas consistent frameworks for animal groups are less developed. Here we suggest a framework for the study of the functional ecology of animals from a trait-based response-effect approach, using dung beetles as model system. Dung beetles are a key group of decomposers that are important for many ecosystem processes. The lack of a trait-based framework tailored to this group has limited the use of traits in dung beetle functional ecology. We review which dung beetle traits respond to the environment and affect ecosystem processes, covering the wide range of spatial, temporal and biological scales at which they are involved. Dung beetles show trait-based responses to variation in temperature, water, soil properties, trophic resources, light, vegetation structure, competition, predation and parasitism. Dung beetles' influence on ecosystem processes includes trait-mediated effects on nutrient cycling, bioturbation, plant growth, seed dispersal, other dung-based organisms and parasite transmission, as well as some cases of pollination and predation. We identify 66 dung beetle traits that are either response or effect traits, or both, pertaining to six main categories: morphology, feeding, reproduction, physiology, activity and movement. Several traits pertain to more than one category, in particular dung relocation behaviour during nesting or feeding. We also identify 136 trait-response and 77 trait-effect relationships in dung beetles. No response to environmental stressors nor effect over ecological processes were related with traits of a single category. This highlights the interrelationship between the traits shaping body-plans, the multi-functionality of traits, and their role linking responses to the environment and effects on the ecosystem. Despite current developments in dung beetle functional ecology, many knowledge gaps remain, and there are biases towards certain traits, functions, taxonomic groups and regions. Our framework provides the foundations for the thorough development of trait-based dung beetle ecology. It also serves as an example framework for other taxa.

Remotely detected aboveground plant function predicts belowground processes in two prairie diversity experiments.

Imaging spectroscopy provides the opportunity to incorporate leaf and canopy optical data into ecological studies, but the extent to which remote sensing of vegetation can enhance the study of belowground processes is not well understood. In terrestrial systems, aboveground and belowground vegetation quantity and quality are coupled, and both influence belowground microbial processes and nutrient cycling. We hypothesized that ecosystem productivity, and the chemical, structural and phylogenetic-functional composition of plant communities would be detectable with remote sensing and could be used to predict belowground plant and soil processes in two grassland biodiversity experiments: the BioDIV experiment at Cedar Creek Ecosystem Science Reserve in Minnesota and the Wood River Nature Conservancy experiment in Nebraska. We tested whether aboveground vegetation chemistry and productivity, as detected from airborne sensors, predict soil properties, microbial processes and community composition. Imaging spectroscopy data were used to map aboveground biomass, green vegetation cover, functional traits and phylogenetic-functional community composition of vegetation. We examined the relationships between the image-derived variables and soil carbon and nitrogen concentration, microbial community composition, biomass and extracellular enzyme activity, and soil processes, including net nitrogen mineralization. In the BioDIV experiment-which has low overall diversity and productivity despite high variation in each-belowground processes were driven mainly by variation in the amount of organic matter inputs to soils. As a consequence, soil respiration, microbial biomass and enzyme activity, and fungal and bacterial composition and diversity were significantly predicted by remotely sensed vegetation cover and biomass. In contrast, at Wood River-where plant diversity and productivity were consistently higher-belowground processes were driven mainly by variation in the quality of aboveground inputs to soils. Consequently, remotely sensed functional, chemical and phylogenetic composition of vegetation predicted belowground extracellular enzyme activity, microbial biomass, and net nitrogen mineralization rates but aboveground biomass (or cover) did not. The contrasting associations between the quantity (productivity) and quality (composition) of aboveground inputs with belowground soil attributes provide a basis for using imaging spectroscopy to understand belowground processes across productivity gradients in grassland systems. However, a mechanistic understanding of how above and belowground components interact among different ecosystems remains critical to extending these results broadly.

Predator Group Composition Indirectly Influences Food Web Dynamics through Predator Growth Rates.

AbstractConsiderable theoretical work predicts that intraspecific trait variation can have profound ecological consequences by altering species interactions. Because of their high flexibility, behavioral traits may be especially relevant in mediating how species respond to one another, thus affecting food web dynamics and ecosystem functioning. However, empirical evidence supporting this idea is limited. Here, we generated predator groups where we manipulated the composition of behavioral types within the groups to assess effects on predator growth rates, prey communities, basal resources, and ecosystem functioning in replicated outdoor ponds. Using European perch (Perca fluviatilis), we created three types of predator populations: two where all individuals expressed either bold or shy phenotypes and one that contained a mix of individuals of the two behavioral types. Bold perch grew faster in mixed populations, indicating that predator growth depends on each individual's behavioral type and that of its group members. However, there was no evidence that the behavioral composition of the perch population directly altered the dynamics of lower trophic levels. Instead, final perch biomass, not behavioral composition, had the strongest influence on lower trophic levels. Thus, the central question may not be whether predator behavior matters at all for trophic dynamics but rather when behavioral effects will predominate over effects of other influences, such as predator biomass variation.

Plant traits controlling growth change in response to a drier climate.

Plant traits are increasingly being used to improve prediction of plant function, including plant demography. However, the capability of plant traits to predict demographic rates remains uncertain, particularly in the context of trees experiencing a changing climate. Here we present data combining 17 plant traits associated with plant structure, metabolism and hydraulic status, with measurements of long-term mean, maximum and relative growth rates for 176 trees from the world's longest running tropical forest drought experiment. We demonstrate that plant traits can predict mean annual tree growth rates with moderate explanatory power. However, only combinations of traits associated more directly with plant functional processes, rather than more commonly employed traits like wood density or leaf mass per area, yield the power to predict growth. Critically, we observe a shift from growth being controlled by traits related to carbon cycling (assimilation and respiration) in well-watered trees, to traits relating to plant hydraulic stress in drought-stressed trees. We also demonstrate that even with a very comprehensive set of plant traits and growth data on large numbers of tropical trees, considerable uncertainty remains in directly interpreting the mechanisms through which traits influence performance in tropical forests.

ForestTemp - Sub-canopy microclimate temperatures of European forests.

Ecological research heavily relies on coarse-gridded climate data based on standardized temperature measurements recorded at 2 m height in open landscapes. However, many organisms experience environmental conditions that differ substantially from those captured by these macroclimatic (i.e. free air) temperature grids. In forests, the tree canopy functions as a thermal insulator and buffers sub-canopy microclimatic conditions, thereby affecting biological and ecological processes. To improve the assessment of climatic conditions and climate-change-related impacts on forest-floor biodiversity and functioning, high-resolution temperature grids reflecting forest microclimates are thus urgently needed. Combining more than 1200 time series of in situ near-surface forest temperature with topographical, biological and macroclimatic variables in a machine learning model, we predicted the mean monthly offset between sub-canopy temperature at 15 cm above the surface and free-air temperature over the period 2000-2020 at a spatial resolution of 25 m across Europe. This offset was used to evaluate the difference between microclimate and macroclimate across space and seasons and finally enabled us to calculate mean annual and monthly temperatures for European forest understories. We found that sub-canopy air temperatures differ substantially from free-air temperatures, being on average 2.1°C (standard deviation ± 1.6°C) lower in summer and 2.0°C higher (±0.7°C) in winter across Europe. Additionally, our high-resolution maps expose considerable microclimatic variation within landscapes, not captured by the gridded macroclimatic products. The provided forest sub-canopy temperature maps will enable future research to model below-canopy biological processes and patterns, as well as species distributions more accurately.

Invertebrate functional traits and terrestrial nutrient cycling: Insights from a global meta-analysis.

Functional traits are useful for characterizing variation in community and ecosystem dynamics. Most advances in trait-based ecology to date centre on plant functional traits, although there is an increasing recognition that animal traits are also key contributors to processes operating at the community or ecosystem scale. Terrestrial invertebrates are incredibly diverse and ubiquitous animals with important roles in nutrient cycling. Despite their widespread influence on ecosystem processes, we currently lack a synthetic understanding of how invertebrate functional traits affect terrestrial nutrient cycling. We present a meta-analysis of 511 paired observations from 122 papers that examined how invertebrate functional traits affected litter decomposition rates, nitrogen pools and litter C:N ratios. Based on the available data, we specifically assessed the effects of feeding mode (bioturbation, detritus shredding, detritus grazing, leaf chewing, leaf piercing, ambush predators, active hunting predators) and body size (macro- and micro-invertebrates) on nutrient cycling. The effects of invertebrates on terrestrial nutrient cycling varied according to functional trait. The inclusion of both macro- (≥2 mm) and micro-invertebrates (<2 mm) increased litter decomposition by 20% and 19%, respectively. All detritivorous feeding modes enhanced litter decomposition rates, with bioturbators, detritus shredders and detritus grazers increasing decomposition by 28%, 22% and 15%, respectively. Neither herbivore feeding mode (e.g. leaf chewers and leaf piercers) nor predator hunting mode (ambush and active hunting) affected decomposition. We also revealed that bioturbators and detritus grazers increased soil nitrogen availability by 99% and 70%, respectively, and that leaf-chewing herbivores had a weak effect on litterfall stoichiometry via reducing C:N ratios by 11%. Although functional traits might be useful predictors of ecosystem processes, our findings suggest context-dependent effects of invertebrate traits on terrestrial nutrient cycling. Detritivore functional traits (i.e. bioturbators, detritus shredders and detritus grazers) are more consistent with increased rates of nutrient cycling, whereas our currently characterized predator and herbivore traits are less predictive. Future research is needed to identify, standardize and deliberately study the impacts of invertebrate functional traits on nutrient cycling in hopes of revealing the key functional traits governing ecosystem functioning worldwide.

Macrophenology: insights into the broad-scale patterns, drivers, and consequences of phenology.

Plant phenology research has surged in recent decades, in part due to interest in phenological sensitivity to climate change and the vital role phenology plays in ecology. Many local-scale studies have generated important findings regarding the physiology, responses, and risks associated with shifts in plant phenology. By comparison, our understanding of regional- and global-scale phenology has been largely limited to remote sensing of green-up without the ability to differentiate among plant species. However, a new generation of analytical tools and data sources-including enhanced remote sensing products, digitized herbarium specimen data, and public participation in science-now permits investigating patterns and drivers of phenology across extensive taxonomic, temporal, and spatial scales, in an emerging field that we call macrophenology. Recent studies have highlighted how phenology affects dynamics at broad scales, including species interactions and ranges, carbon fluxes, and climate. At the cusp of this developing field of study, we review the theoretical and practical advances in four primary areas of plant macrophenology: (1) global patterns and shifts in plant phenology, (2) within-species changes in phenology as they mediate species' range limits and invasions at the regional scale, (3) broad-scale variation in phenology among species leading to ecological mismatches, and (4) interactions between phenology and global ecosystem processes. To stimulate future research, we describe opportunities for macrophenology to address grand challenges in each of these research areas, as well as recently available data sources that enhance and enable macrophenology research.

Foliar functional traits from imaging spectroscopy across biomes in eastern North America.

Foliar functional traits are widely used to characterize leaf and canopy properties that drive ecosystem processes and to infer physiological processes in Earth system models. Imaging spectroscopy provides great potential to map foliar traits to characterize continuous functional variation and diversity, but few studies have demonstrated consistent methods for mapping multiple traits across biomes. With airborne imaging spectroscopy data and field data from 19 sites, we developed trait models using partial least squares regression, and mapped 26 foliar traits in seven NEON (National Ecological Observatory Network) ecoregions (domains) including temperate and subtropical forests and grasslands of eastern North America. Model validation accuracy varied among traits (normalized root mean squared error, 9.1-19.4%; coefficient of determination, 0.28-0.82), with phenolic concentration, leaf mass per area and equivalent water thickness performing best across domains. Across all trait maps, 90% of vegetated pixels had reasonable values for one trait, and 28-81% provided high confidence for multiple traits concurrently. Maps of 26 traits and their uncertainties for eastern US NEON sites are available for download, and are being expanded to the western United States and tundra/boreal zone. These data enable better understanding of trait variations and relationships over large areas, calibration of ecosystem models, and assessment of continental-scale functional diversity.

SoilTemp: A global database of near-surface temperature.

Current analyses and predictions of spatially explicit patterns and processes in ecology most often rely on climate data interpolated from standardized weather stations. This interpolated climate data represents long-term average thermal conditions at coarse spatial resolutions only. Hence, many climate-forcing factors that operate at fine spatiotemporal resolutions are overlooked. This is particularly important in relation to effects of observation height (e.g. vegetation, snow and soil characteristics) and in habitats varying in their exposure to radiation, moisture and wind (e.g. topography, radiative forcing or cold-air pooling). Since organisms living close to the ground relate more strongly to these microclimatic conditions than to free-air temperatures, microclimatic ground and near-surface data are needed to provide realistic forecasts of the fate of such organisms under anthropogenic climate change, as well as of the functioning of the ecosystems they live in. To fill this critical gap, we highlight a call for temperature time series submissions to SoilTemp, a geospatial database initiative compiling soil and near-surface temperature data from all over the world. Currently, this database contains time series from 7,538 temperature sensors from 51 countries across all key biomes. The database will pave the way toward an improved global understanding of microclimate and bridge the gap between the available climate data and the climate at fine spatiotemporal resolutions relevant to most organisms and ecosystem processes.

Habitat patchiness, ecological connectivity and the uneven recovery of boreal stream ecosystems from an experimental drought.

Ongoing climate change is increasing the occurrence and intensity of drought episodes worldwide, including in boreal regions not previously regarded as drought prone, and where the impacts of drought remain poorly understood. Ecological connectivity is one factor that might influence community structure and ecosystem functioning post-drought, by facilitating the recovery of sensitive species via dispersal at both local (e.g. a nearby habitat patch) and regional (from other systems within the same region) scales. In an outdoor mesocosm experiment, we investigated how impacts of drought on boreal stream ecosystems are altered by the spatial arrangement of local habitat patches within stream channels, and variation in ecological connectivity with a regional species pool. We measured basal ecosystem processes underlying carbon and nutrient cycling: (a) algal biomass accrual; (b) microbial respiration; and (c) decomposition of organic matter, and sampled communities of aquatic fungi and benthic invertebrates. An 8-day drought event had strong impacts on both community structure and ecosystem functioning, including algal accrual, leaf decomposition and microbial respiration, with many of these impacts persisting even after water levels had been restored for 3.5 weeks. Enhanced connectivity with the regional species pool and increased aggregation of habitat patches also affected multiple response variables, especially those associated with microbes, and in some cases reduced the effects of drought to a small extent. This indicates that spatial processes might play a role in the resilience of communities and ecosystem functioning, given enough time. These effects were however insufficient to facilitate significant recovery in algal growth before seasonal dieback began in autumn. The limited resilience of ecosystem functioning in our experiment suggests that even short-term droughts can have extended consequences for stream ecosystems in the world's vast boreal region, and especially on the ecosystem processes and services mediated by algal biofilms.

Investigating changes in estuarine ecosystem functioning under future scenarios.

Estuaries are subject to disturbance by land-based sediment and nutrient inputs, resulting in changes to the ecosystems and the functions and services that they support. Spatial mapping tools that identify how functional hotspots in the estuary may shift in location and intensity under different disturbance scenarios highlight to managers the trajectory of change and the value of active management and restoration, but to date these tools are only available in the most intensively researched ecosystems. Using empirical data derived from long-term monitoring and multi-habitat field experiments we developed future scenarios representing different impacts of environmental degradation on estuarine ecosystem functions that are important for supporting ecosystem services. We used the spatial prioritization software Zonation in a novel fashion to assess effects of different disturbance scenarios on critical soft-sediment ecosystem processes (nutrient fluxes and sediment erodibility measures) that are influenced by macrofaunal communities and local environment conditions. We compared estimates of current conditions with three scenarios linked to changes in land-use and resulting downstream impacts on estuarine ecosystems to determine how disturbance influences the distribution of high value areas for ecosystem function. Scenarios investigated the implications of habitat degradation associated with sediment deposition and declines in large sediment-dwelling animal abundance whose behavior has important influences on ecosystem function. Our analyses demonstrate decreases in the majority of ecosystem processes under scenarios associated with disturbances. These results suggest that it is important to restore biodiversity and ecosystem function and that the application of Zonation in this context offers a simple, rapid and cost-effective way of identifying priority actions and locations for restoration, and how these shift due to multiple impacts.

Below-ground frontiers in trait-based plant ecology.

Contents 1597 I. 1597 II. 1597 III. 1598 IV. 1598 V. 1600 VI. 1601 VII. 1601 VIII. 1601 1602 References 1602 SUMMARY: Trait-based approaches have led to significant advances in plant ecology, but are currently biased toward above-ground traits. It is becoming clear that a stronger emphasis on below-ground traits is needed to better predict future changes in plant biodiversity and their consequences for ecosystem functioning. Here I propose six 'below-ground frontiers' in trait-based plant ecology, with an emphasis on traits governing soil nutrient acquisition: redefining fine roots; quantifying root trait dimensionality; integrating mycorrhizas; broadening the suite of root traits; determining linkages between root traits and abiotic and biotic factors; and understanding ecosystem-level consequences of root traits. Focusing research efforts along these frontiers should help to fulfil the promise of trait-based ecology: enhanced predictive capacity across ecological scales.

The changing contribution of top-down and bottom-up limitation of mesopredators during 220 years of land use and climate change.

Apex predators may buffer bottom-up driven ecosystem change, as top-down suppression may dampen herbivore and mesopredator responses to increased resource availability. However, theory suggests that for this buffering capacity to be realized, the equilibrium abundance of apex predators must increase. This raises the question: will apex predators maintain herbivore/mesopredator limitation, if bottom-up change relaxes resource constraints? Here, we explore changes in mesopredator (red fox Vulpes vulpes) abundance over 220 years in response to eradication and recovery of an apex predator (Eurasian lynx Lynx lynx), and changes in land use and climate which are linked to resource availability. A three-step approach was used. First, recent data from Finland and Sweden were modelled to estimate linear effects of lynx density, land use and winter temperature on fox density. Second, lynx density, land use and winter temperature was estimated in a 22 650 km2 focal area in boreal and boreo-nemoral Sweden in the years 1830, 1920, 2010 and 2050. Third, the models and estimates were used to project historic and future fox densities in the focal area. Projected fox density was lowest in 1830 when lynx density was high, winters cold and the proportion of cropland low. Fox density peaked in 1920 due to lynx eradication, a mesopredator release boosted by favourable bottom-up changes - milder winters and cropland expansion. By 2010, lynx recolonization had reduced fox density, but it remained higher than in 1830, partly due to the bottom-up changes. Comparing 1830 to 2010, the contribution of top-down limitation decreased, while environment enrichment relaxed bottom-up limitation. Future scenarios indicated that by 2050, lynx density would have to increase by 79% to compensate for a projected climate-driven increase in fox density. We highlight that although top-down limitation in theory can buffer bottom-up change, this requires compensatory changes in apex predator abundance. Hence apex predator recolonization/recovery to historical levels would not be sufficient to compensate for widespread changes in climate and land use, which have relaxed the resource constraints for many herbivores and mesopredators. Variation in bottom-up conditions may also contribute to context dependence in apex predator effects.

What determines the importance of a species for ecosystem processes? Insights from tropical ant assemblages.

Biodiversity is known to increase ecosystem functioning. However, species vary in their contributions to ecosystem processes. Here, we investigated seven ecosystem functions based on the consumption of different resources in tropical ant communities. We analysed how different species influence site-level resource consumption, and determined how each species influenced performance and stability of these functions. Based on simulated extinctions, we identified 'key species' with significant functional contributions. We then investigated which traits, such as biomass, abundance, and specialisation, characterized them, and compared trait distributions across four sites to analyse differences in functional redundancy. Only few species significantly influenced ecosystem functions. Common generalist species tended to be the most important drivers of many ecosystem functions, though several specialist species also proved to be important in this study. Moreover, species-specific ecological impacts varied across sites. In addition, we found that functional redundancy varied across sites, and was highest in sites where the most common species did not simultaneously have the greatest functional impacts. Furthermore, redundancy was enhanced in sites where species were less specialised and had more even incidence distributions. Our study demonstrates that the ecological importance of a species depends on its functional traits, but also on the community context. It cannot be assessed without investigating its species-specific performance across multiple functions. Hence, to assess functional redundancy in a habitat and the potential for compensation of species loss, researchers need to study species-specific traits that concern functional performance as well as population dynamics and tolerance to environmental conditions.