Integrating natural gradients, experiments, and statistical modeling in a distributed network experiment: An example from the WaRM Network

A growing body of work examines the direct and indirect effects of climate change on ecosystems, typically by using manipulative experiments at a single site or performing meta-analyses across many independent experiments. However, results from single-site studies tend to have limited generality. Although meta-analytic approaches can help overcome this by exploring trends across sites, the inherent limitations in combining disparate datasets from independent approaches remain a major challenge. In this paper, we present a globally distributed experimental network that can be used to disentangle the direct and indirect effects of climate change. We discuss how natural gradients, experimental approaches, and statistical techniques can be combined to best inform predictions about responses to climate change, and we present a globally distributed experiment that utilizes natural environmental gradients to better understand long-term community and ecosystem responses to environmental change. The warming and (species) removal in mountains (WaRM) network employs experimental warming and plant species removals at high- and low-elevation sites in a factorial design to examine the combined and relative effects of climatic warming and the loss of dominant species on community structure and ecosystem function, both above- and belowground. The experimental design of the network allows for increasingly common statistical approaches to further elucidate the direct and indirect effects of warming. We argue that combining ecological observations and experiments along gradients is a powerful approach to make stronger predictions of how ecosystems will function in a warming world as species are lost, or gained, in local communities

Integrating natural gradients, experiments, and statistical modeling in a distributed network experiment: An example from the WaRM Network

A growing body of work examines the direct and indirect effects of climate change on ecosystems, typically by using manipulative experiments at a single site or performing meta-analyses across many independent experiments. However, results from single-site studies tend to have limited generality. Although meta-analytic approaches can help overcome this by exploring trends across sites, the inherent limitations in combining disparate datasets from independent approaches remain a major challenge. In this paper, we present a globally distributed experimental network that can be used to disentangle the direct and indirect effects of climate change. We discuss how natural gradients, experimental approaches, and statistical techniques can be combined to best inform predictions about responses to climate change, and we present a globally distributed experiment that utilizes natural environmental gradients to better understand long-term community and ecosystem responses to environmental change. The warming and (species) removal in mountains (WaRM) network employs experimental warming and plant species removals at high- and low-elevation sites in a factorial design to examine the combined and relative effects of climatic warming and the loss of dominant species on community structure and ecosystem function, both above- and belowground. The experimental design of the network allows for increasingly common statistical approaches to further elucidate the direct and indirect effects of warming. We argue that combining ecological observations and experiments along gradients is a powerful approach to make stronger predictions of how ecosystems will function in a warming world as species are lost, or gained, in local communities

Circum-Arctic distribution of chemical anti-herbivore compounds suggests biome-wide trade-off in defence strategies in Arctic shrubs

Spatial variation in plant chemical defence towards herbivores can help us understand variation in herbivore top-down control of shrubs in the Arctic and possibly also shrub responses to global warming. Less defended, non-resinous shrubs could be more influenced by herbivores than more defended, resinous shrubs. However, sparse field measurements limit our current understanding of how much of the circum-Arctic variation in defence compounds is explained by taxa or defence functional groups (resinous/non-resinous). We measured circum-Arctic chemical defence and leaf digestibility in resinous (Betula glandulosa, B. nana ssp. exilis) and non-resinous (B. nana ssp. nana, B. pumila) shrub birches to see how they vary among and within taxa and functional groups. Using liquid chromatography-mass spectrometry (LC-MS) metabolomic analyses and in vitro leaf digestibility via incubation in cattle rumen fluid, we analysed defence composition and leaf digestibility in 128 samples from 44 tundra locations. We found biogeographical patterns in anti-herbivore defence where mean leaf triterpene concentrations and twig resin gland density were greater in resinous taxa and mean concentrations of condensing tannins were greater in non-resinous taxa. This indicates a biome-wide trade-off between triterpene- or tannin-dominated defences. However, we also found variations in chemical defence composition and resin gland density both within and among functional groups (resinous/non-resinous) and taxa, suggesting these categorisations only partly predict chemical herbivore defence. Complex tannins were the only defence compounds negatively related to in vitro digestibility, identifying this previously neglected tannin group as having a potential key role in birch anti-herbivore defence. We conclude that circum-Arctic variation in birch anti-herbivore defence can be partly derived from biogeographical distributions of birch taxa, although our detailed mapping of plant defence provides more information on this variation and can be used for better predictions of herbivore effects on Arctic vegetation.Peer reviewe

Circum-Arctic distribution of chemical anti-herbivore compounds suggests biome-wide trade-off in defence strategies in Arctic shrubs

Spatial variation in plant chemical defence towards herbivores can help us understand variation in herbivore top?down control of shrubs in the Arctic and possibly also shrub responses to global warming. Less defended, non-resinous shrubs could be more influenced by herbivores than more defended, resinous shrubs. However, sparse field measurements limit our current understanding of how much of the circum-Arctic variation in defence compounds is explained by taxa or defence functional groups (resinous/non-resinous). We measured circum-Arctic chemical defence and leaf digestibility in resinous (Betula glandulosa, B. nana ssp. exilis) and non-resinous (B. nana ssp. nana, B. pumila) shrub birches to see how they vary among and within taxa and functional groups. Using liquid chromatography?mass spectrometry (LC?MS) metabolomic analyses and in vitro leaf digestibility via incubation in cattle rumen fluid, we analysed defence composition and leaf digestibility in 128 samples from 44 tundra locations. We found biogeographical patterns in anti-herbivore defence where mean leaf triterpene concentrations and twig resin gland density were greater in resinous taxa and mean concentrations of condensing tannins were greater in non-resinous taxa. This indicates a biome-wide trade-off between triterpene- or tannin-dominated defences. However, we also found variations in chemical defence composition and resin gland density both within and among functional groups (resinous/non-resinous) and taxa, suggesting these categorisations only partly predict chemical herbivore defence. Complex tannins were the only defence compounds negatively related to in vitro digestibility, identifying this previously neglected tannin group as having a potential key role in birch anti-herbivore defence. We conclude that circum-Arctic variation in birch anti-herbivore defence can be partly derived from biogeographical distributions of birch taxa, although our detailed mapping of plant defence provides more information on this variation and can be used for better predictions of herbivore effects on Arctic vegetation

Plant and microbial responses to nitrogen and phosphorus addition across an elevational gradient in subarctic tundra

Temperature and nutrients are major limiting factors in subarctic tundra. Experimental manipulation of nutrient availability along elevational gradients (and thus temperature) can improve our understanding of ecological responses to climate change. However, no study to date has explored impacts of nutrient addition along a tundra elevational gradient, or across contrasting vegetation types along any elevational gradient. We set up a full factorial nitrogen (N) and phosphorus (P) fertilization experiment in each of two vegetation types (heath and meadow) at 500 m, 800 m, and 1000 m elevation in northern Swedish tundra. We predicted that plant and microbial communities in heath or at lower elevations would be more responsive to N addition while communities in meadow or at higher elevations would be more responsive to P addition, and that fertilizer effects would vary more with elevation for the heath than for the meadow. Although our results provided little support for these predictions, the relationship between nutrient limitation and elevation differed between vegetation types. Most plant and microbial properties were responsive to N and/or P fertilization, but responses often varied with elevation and/or vegetation type. For instance, vegetation density significantly increased with N + P fertilization relative to the other fertilizer treatments, and this increase was greatest at the lowest elevation for the heath but at the highest elevation for the meadow. Arbuscular mycorrhizae decreased with P fertilization at 500 m for the meadow, but with all fertilizer treatments in both vegetation types at 800 m. Fungal to bacterial ratios were enhanced by N + P fertilization for the two highest elevations in the meadow only. Additionally, microbial responses to fertilization were primarily direct rather than indirect via plant responses, pointing to a decoupled response of plant and microbial communities to nutrient addition and elevation. Because our study shows how two community types differ in their responses to fertilization and elevation, and because the temperature range across this gradient is similar to 3 degrees C, our study is informative about how nutrient limitation in tundra may be influenced by temperature shifts that are comparable to those expected under climate change during this century

Data from: Environmental factors and traits that drive plant litter decomposition do not determine home-field advantage effects

The ‘home-field advantage’ (HFA) hypothesis predicts that plant litter is decomposed faster than expected underneath the plant from which it originates (‘home’) than underneath other plants (‘away’), because decomposer communities are specialized to break down litter from the plants they associate with. However, empirical evidence shows that the occurrence of HFA is highly variable, and the reasons for this are little understood. In our study we progress our understanding by investigating whether HFA is stronger for more recalcitrant litter types and under colder conditions and how soil properties and plant functional traits affect the magnitude and direction of HFA. In subarctic tundra in northern Sweden we set up a reciprocal transplant litter decomposition experiment along an elevational gradient where three highly contrasting vegetation types (heath, meadow and Salix) occur at all elevations, and where temperature decreases strongly with elevation. In this study, we used a litter bag approach where litters from each elevation × vegetation type combination were decomposed in all combinations of elevation × vegetation type. We also measured community-level plant functional traits, such as leaf and litter nutrient content. We determined soil biotic and abiotic properties, such as microbial biomass and soil nutrient content, in soil cores collected for each elevation × vegetation type combination. We found that mass loss increased with plant and litter nutrient content and with soil temperature. In contrast, the occurrence of HFA was limited in our study system, and its magnitude and direction could not be explained by vegetation type, elevation, plant traits or soil properties, despite these factors serving as powerful drivers of litter mass loss in our study. We conclude that although vegetation type and climate are major drivers of litter mass loss, they do not emerge as important determinants of HFA. Therefore, while rapid shifts in plant community composition or temperature due to global change are likely to influence litter mass loss directly by altering environmental conditions, plant trait spectra and litter quality, indirect effects of global change resulting from decoupling of specialist interactions between litter and decomposer communities appears to be of less importance

Plant and microbial responses to nitrogen and phosphorus addition across an elevational gradient in subarctic tundra

Temperature and nutrients are major limiting factors in subarctic tundra. Experimental manipulation of nutrient availability along elevational gradients (and thus temperature) can improve our understanding of ecological responses to climate change. However, no study to date has explored impacts of nutrient addition along a tundra elevational gradient, or across contrasting vegetation types along any elevational gradient. We set up a full factorial nitrogen (N) and phosphorus (P) fertilization experiment in each of two vegetation types (heath and meadow) at 500 m, 800 m, and 1000 m elevation in northern Swedish tundra. We predicted that plant and microbial communities in heath or at lower elevations would be more responsive to N addition while communities in meadow or at higher elevations would be more responsive to P addition, and that fertilizer effects would vary more with elevation for the heath than for the meadow. Although our results provided little support for these predictions, the relationship between nutrient limitation and elevation differed between vegetation types. Most plant and microbial properties were responsive to N and/or P fertilization, but responses often varied with elevation and/or vegetation type. For instance, vegetation density significantly increased with N + P fertilization relative to the other fertilizer treatments, and this increase was greatest at the lowest elevation for the heath but at the highest elevation for the meadow. Arbuscular mycorrhizae decreased with P fertilization at 500 m for the meadow, but with all fertilizer treatments in both vegetation types at 800 m. Fungal to bacterial ratios were enhanced by N + P fertilization for the two highest elevations in the meadow only. Additionally, microbial responses to fertilization were primarily direct rather than indirect via plant responses, pointing to a decoupled response of plant and microbial communities to nutrient addition and elevation. Because our study shows how two community types differ in their responses to fertilization and elevation, and because the temperature range across this gradient is similar to 3 degrees C, our study is informative about how nutrient limitation in tundra may be influenced by temperature shifts that are comparable to those expected under climate change during this century

Uneven global distribution of food web studies under climate change

Abstract Trophic interactions within food webs affect species distributions, coexistence, and provision of ecosystem services but can be strongly impacted by climatic changes. Understanding these impacts is therefore essential for managing ecosystems and sustaining human well-being. Here, we conducted a global synthesis of terrestrial, marine, and freshwater studies to identify key gaps in our knowledge of climate change impacts on food webs and determine whether the areas currently studied are those most likely to be impacted by climate change. We found research suffers from a strong geographic bias, with only 3.5% of studies occurring in the tropics. Importantly, the distribution of sites sampled under projected climate changes was biased?areas with decreases or large increases in precipitation and areas with low magnitudes of temperature change were under-represented. Our results suggest that understanding of climate change impacts on food webs could be broadened by considering more than two trophic levels, responses in addition to species abundance and biomass, impacts of a wider suite of climatic variables, and tropical ecosystems. Most importantly, to enable better forecasts of biodiversity responses to climate change, we identify critically under-represented geographic regions and climatic conditions which should be prioritized in future research.Peer reviewe

Decomposition rate and stabilization across six tundra vegetation types exposed to >20 years of warming

Aims: Litter decomposition is an important driver of soil carbon and nutrient cycling in nutrient-limited Arctic ecosystems. However, climate change is expected to induce changes that directly or indirectly affect decomposition. We examined the direct effects of long-term warming relative to differences in soil abiotic properties associated with vegetation type on litter decomposition across six subarctic vegetation types. Methods: In six vegetation types, rooibos and green tea bags were buried for 70–75 days at 8 cm depth inside warmed (by open-top chambers) and control plots that had been in place for 20–25 years. Standardized initial decomposition rate and stabilization of the labile material fraction of tea (into less decomposable material) were calculated from tea mass losses. Soil moisture and temperature were measured bi-weekly during summer and plant-available nutrients were measured with resin probes. Results: Initial decomposition rate was decreased by the warming treatment. Stabilization was less affected by warming and determined by vegetation type and soil moisture. Soil metal concentrations impeded both initial decomposition rate and stabilization. Conclusions: While a warmer Arctic climate will likely have direct effects on initial litter decomposition rates in tundra, stabilization of organic matter was more affected by vegetation type and soil parameters and less prone to be affected by direct effects of warming