Soil nematode abundance and functional group composition at a global scale.

Soil organisms are a crucial part of the terrestrial biosphere. Despite their importance for ecosystem functioning, few quantitative, spatially explicit models of the active belowground community currently exist. In particular, nematodes are the most abundant animals on Earth, filling all trophic levels in the soil food web. Here we use 6,759 georeferenced samples to generate a mechanistic understanding of the patterns of the global abundance of nematodes in the soil and the composition of their functional groups. The resulting maps show that 4.4 ± 0.64 × 1020 nematodes (with a total biomass of approximately 0.3 gigatonnes) inhabit surface soils across the world, with higher abundances in sub-Arctic regions (38% of total) than in temperate (24%) or tropical (21%) regions. Regional variations in these global trends also provide insights into local patterns of soil fertility and functioning. These high-resolution models provide the first steps towards representing soil ecological processes in global biogeochemical models and will enable the prediction of elemental cycling under current and future climate scenarios.

The biological controls of soil carbon accumulation following wildfire and harvest in boreal forests: A review.

Boreal forests are frequently subjected to disturbances, including wildfire and clear-cutting. While these disturbances can cause soil carbon (C) losses, the long-term accumulation dynamics of soil C stocks during subsequent stand development is controlled by biological processes related to the balance of net primary production (NPP) and outputs via heterotrophic respiration and leaching, many of which remain poorly understood. We review the biological processes suggested to influence soil C accumulation in boreal forests. Our review indicates that median C accumulation rates following wildfire and clear-cutting are similar (0.15 and 0.20 Mg ha-1 year-1, respectively), however, variation between studies is extremely high. Further, while many individual studies show linear increases in soil C stocks through time after disturbance, there are indications that C stock recovery is fastest early to mid-succession (e.g. 15-80 years) and then slows as forests mature (e.g. >100 years). We indicate that the rapid build-up of soil C in younger stands appears not only driven by higher plant production, but also by a high rate of mycorrhizal hyphal production, and mycorrhizal suppression of saprotrophs. As stands mature, the balance between reductions in plant and mycorrhizal production, increasing plant litter recalcitrance, and ectomycorrhizal decomposers and saprotrophs have been highlighted as key controls on soil C accumulation rates. While some of these controls appear well understood (e.g. temporal patterns in NPP, changes in aboveground litter quality), many others remain research frontiers. Notably, very little data exists describing and comparing successional patterns of root production, mycorrhizal functional traits, mycorrhizal-saprotroph interactions, or C outputs via heterotrophic respiration and dissolved organic C following different disturbances. We argue that these less frequently described controls require attention, as they will be key not only for understanding ecosystem C balances, but also for representing these dynamics more accurately in soil organic C and Earth system models.

Isopod mouthpart traits respond to a tropical forest recovery gradient.

Functional trait ecology has the potential to provide generalizable and mechanistic predictions of ecosystem function from data of species distributions and traits. The traits that are selected should both respond to environmental factors and influence ecosystem functioning. Invertebrate mouthpart traits fulfill these criteria, but are seldom collected, lack standardized measurement protocols, and have infrequently been investigated in response to environmental factors. We surveyed isopod species that consume plant detritus, and tree communities in 58 plots across primary and secondary forests in Singapore. We measured body dimensions (body size traits), pereopod and antennae lengths (locomotory traits), dimensions of mandible structures (morphological mouthpart traits), and mechanical advantages generated by mandible shape (mechanical mouthpart traits) for six isopod species found in these plots and investigated if these traits respond to changes in tree community composition, tree diversity, and forest structure. Morphological mouthpart traits responded to a tree compositional gradient reflecting forest recovery degree. Mouthpart features associated with greater consumption of litter (broader but less serrated/rugose lacinia mobilis [an important cutting and chewing structure on the mandible]) were most prevalent in abandoned plantation and young secondary forests containing disturbance-associated tree species. Feeding strategies associated with fungi grazing (narrower and more serrated/rugose lacinia mobilis) were most prevalent in late secondary forests containing later successional tree species. Since morphological mouthpart traits likely also predict consumption and excretion rates of isopods, these traits advance our understanding of environment-trait-ecosystem functioning relationships across contrasting tropical forest plots that vary in composition, disturbance history, and post-disturbance recovery.

Long-term effects of species loss on community properties across contrasting ecosystems.

Biodiversity loss can heavily affect the functioning of ecosystems, and improving our understanding of how ecosystems respond to biodiversity decline is one of the main challenges in ecology1-4. Several important aspects of the longer-term effects of biodiversity loss on ecosystems remain unresolved, including how these effects depend on environmental context5-7. Here we analyse data from an across-ecosystem biodiversity manipulation experiment that, to our knowledge, represents the world's longest-running experiment of this type. This experiment has been set up on 30 lake islands in Sweden that vary considerably in productivity and soil fertility owing to differences in fire history8,9. We tested the effects of environmental context on how plant species loss affected two fundamental community attributes-plant community biomass and temporal variability-over 20 years. In contrast to findings from artificially assembled communities10-12, we found that the effects of species loss on community biomass decreased over time; this decrease was strongest on the least productive and least fertile islands. Species loss generally also increased temporal variability, and these effects were greatest on the most productive and most fertile islands. Our findings highlight that the ecosystem-level consequences of biodiversity loss are not constant across ecosystems and that understanding and forecasting these consequences necessitates taking into account the overarching role of environmental context.

From bottom-up to top-down control of invertebrate herbivores in a retrogressive chronosequence.

In the long-term absence of disturbance, ecosystems often enter a decline or retrogressive phase which leads to reductions in primary productivity, plant biomass, nutrient cycling and foliar quality. However, the consequences of ecosystem retrogression for higher trophic levels such as herbivores and predators, are less clear. Using a post-fire forested island-chronosequence across which retrogression occurs, we provide evidence that nutrient availability strongly controls invertebrate herbivore biomass when predators are few, but that there is a switch from bottom-up to top-down control when predators are common. This trophic flip in herbivore control probably arises because invertebrate predators respond to alternative energy channels from the adjacent aquatic matrix, which were independent of terrestrial plant biomass. Our results suggest that effects of nutrient limitation resulting from ecosystem retrogression on trophic cascades are modified by nutrient-independent variation in predator abundance, and this calls for a more holistic approach to trophic ecology to better understand herbivore effects on plant communities.

Leaf decomposition and flammability are largely decoupled across species in a tropical swamp forest despite sharing some predictive leaf functional traits.

Decomposition and fire are major carbon pathways in many ecosystems, yet potential linkages between these processes are poorly understood. We test whether variability in decomposability and flammability across species are related to each other and to key plant functional traits in tropical swamp forests, where habitat degradation is elevating decomposition and fire regimes. Using senesced and fresh leaves of 22 swamp tree species in Singapore, we conducted an in situ decomposition experiment and a laboratory flammability experiment. We analysed 16 leaf physical and biochemical traits as predictors of decomposability and components of flammability: combustibility, ignitability and sustainability. Decomposability and flammability were largely decoupled across species, despite some shared predictive traits such as specific leaf area (SLA). Physical traits predicted that thicker leaves with a smaller SLA and volume decomposed faster, while various cation concentrations predicted flammability components, particularly ignitability. We show that flammability and decomposability of swamp forest leaves are decoupled because flammability is mostly driven by biochemical traits, while decomposition is driven by physical traits. Our approach identifies species that are slow to decompose and burn (e.g. Calophyllum tetrapterum and Xanthophyllum flavescens), which could be planted to mitigate carbon losses in tropical swamp reforestation.

Temporal dynamics of soil fungi in a pyrodiverse dry-sclerophyll forest.

Fire is a major evolutionary and ecological driver that shapes biodiversity in forests. While above-ground community responses to fire have been well-documented, those below-ground are much less understood. However, below-ground communities, including fungi, play key roles in forests and facilitate the recovery of other organisms after fire. Here, we used internal transcribed spacer (ITS) meta-barcoding data from forests with three different times since fire [short (3 years), medium (13-19 years) and long (>26 years)] to characterize the temporal responses of soil fungal communities across functional groups, ectomycorrhizal exploration strategies and inter-guild associations. Our findings indicate that fire effects on fungal communities are strongest in the short to medium term, with clear distinctions between communities in forests with a short time (3 years) since fire, a medium time (13-19 years) and a long time (>26 years) since fire. Ectomycorrhizal fungi were disproportionately impacted by fire relative to saprotrophs, but the direction of the response varied depending on morphological structures and exploration strategies. For instance, short-distance ectomycorrhizal fungi increased with recent fire, while medium-distance (fringe) ectomycorrhizal fungi decreased. Further, we detected strong, negative inter-guild associations between ectomycorrhizal and saprotrophic fungi but only at medium and long times since fire. Given the functional significance of fungi, the temporal changes in fungal composition, inter-guild associations and functional groups after fire demonstrated in our study may have functional implications that require adaptive management to curtail.

Linking the influence and dependence of people on biodiversity across scales.

Biodiversity enhances many of nature's benefits to people, including the regulation of climate and the production of wood in forests, livestock forage in grasslands and fish in aquatic ecosystems. Yet people are now driving the sixth mass extinction event in Earth's history. Human dependence and influence on biodiversity have mainly been studied separately and at contrasting scales of space and time, but new multiscale knowledge is beginning to link these relationships. Biodiversity loss substantially diminishes several ecosystem services by altering ecosystem functioning and stability, especially at the large temporal and spatial scales that are most relevant for policy and conservation.

Elevation alters ecosystem properties across temperate treelines globally.

Temperature is a primary driver of the distribution of biodiversity as well as of ecosystem boundaries. Declining temperature with increasing elevation in montane systems has long been recognized as a major factor shaping plant community biodiversity, metabolic processes, and ecosystem dynamics. Elevational gradients, as thermoclines, also enable prediction of long-term ecological responses to climate warming. One of the most striking manifestations of increasing elevation is the abrupt transitions from forest to treeless alpine tundra. However, whether there are globally consistent above- and belowground responses to these transitions remains an open question. To disentangle the direct and indirect effects of temperature on ecosystem properties, here we evaluate replicate treeline ecotones in seven temperate regions of the world. We find that declining temperatures with increasing elevation did not affect tree leaf nutrient concentrations, but did reduce ground-layer community-weighted plant nitrogen, leading to the strong stoichiometric convergence of ground-layer plant community nitrogen to phosphorus ratios across all regions. Further, elevation-driven changes in plant nutrients were associated with changes in soil organic matter content and quality (carbon to nitrogen ratios) and microbial properties. Combined, our identification of direct and indirect temperature controls over plant communities and soil properties in seven contrasting regions suggests that future warming may disrupt the functional properties of montane ecosystems, particularly where plant community reorganization outpaces treeline advance.

Ericoid shrubs shape fungal communities and suppress organic matter decomposition in boreal forests.

Mycorrhizal fungi associated with boreal trees and ericaceous shrubs are central actors in organic matter (OM) accumulation through their belowground carbon allocation, their potential capacity to mine organic matter for nitrogen (N) and their ability to suppress saprotrophs. Yet, interactions between co-occurring ectomycorrhizal fungi (EMF), ericoid mycorrhizal fungi (ERI), and saprotrophs are poorly understood. We used a long-term (19 yr) plant functional group manipulation experiment with removals of tree roots, ericaceous shrubs and mosses and analysed the responses of different fungal guilds (assessed by metabarcoding) and their interactions in relation to OM quality (assessed by mid-infrared spectroscopy and nuclear magnetic resonance) and decomposition (litter mesh-bags) across a 5000-yr post-fire boreal forest chronosequence. We found that the removal of ericaceous shrubs and associated ERI changed the composition of EMF communities, with larger effects occurring at earlier stages of the chronosequence. Removal of shrubs was associated with enhanced N availability, litter decomposition and enrichment of the recalcitrant OM fraction. We conclude that increasing abundance of slow-growing ericaceous shrubs and the associated fungi contributes to increasing nutrient limitation, impaired decomposition and progressive OM accumulation in boreal forests, particularly towards later successional stages. These results are indicative of the contrasting roles of EMF and ERI in regulating belowground OM storage.

Responses of bryosphere fauna to drought across a boreal forest chronosequence.

Projected changes in precipitation regimes can greatly impact soil biota, which in turn alters key ecosystem functions. In moss-dominated ecosystems, the bryosphere (i.e., the ground moss layer including live and senesced moss) plays a key role in carbon and nutrient cycling, and it hosts high abundances of microfauna (i.e., nematodes and tardigrades) and mesofauna (i.e., mites and springtails). However, we know very little about how bryosphere fauna responds to precipitation, and whether this response changes across environmental gradients. Here, we used a mesocosm experiment to study the effect of volume and frequency of precipitation on the abundance and community composition of functional groups of bryosphere fauna. Hylocomium splendens bryospheres were sampled from a long-term post-fire boreal forest chronosequence in northern Sweden which varies greatly in environmental conditions. We found that reduced precipitation promoted the abundance of total microfauna and of total mesofauna, but impaired predaceous/omnivorous nematodes, and springtails. Generally, bryosphere fauna responded more strongly to precipitation volume than to precipitation frequency. For some faunal functional groups, the effects of precipitation frequency were stronger at reduced precipitation volumes. Context-dependency effects were found for microfauna only: microfauna was more sensitive to precipitation in late-successional forests (i.e., those with lower productivity and soil nutrient availability) than in earlier-successional forests. Our results also suggest that drought-induced changes in trophic interactions and food resources in the bryosphere may increase faunal abundance. Consequently, drier bryospheres that may result from climate change could promote carbon and nutrient turnover from fauna activity, especially in older, less productive forests.

Home-field advantage of litter decomposition: from the phyllosphere to the soil.

Plants often associate with specialized decomposer communities that increase plant litter breakdown, a phenomenon that is known as the 'home-field advantage' (HFA). Although the concept of HFA has long considered only the role of the soil microbial community, explicit consideration of the role of the microbial community on the foliage before litter fall (i.e. the phyllosphere community) may help us to better understand HFA. We investigated the occurrence of HFA in the presence vs absence of phyllosphere communities and found that HFA effects were smaller when phyllosphere communities were removed. We propose that priority effects and interactions between phyllosphere and soil organisms can help explain the positive effects of the phyllosphere at home, and suggest a path forward for further investigation.

Root trait-microbial relationships across tundra plant species.

Fine roots, and their functional traits, influence associated rhizosphere microorganisms via root exudation and root litter quality. However, little information is known about their relationship with rhizosphere microbial taxa and functional guilds. We investigated the relationships of 11 fine root traits of 20 sub-arctic tundra meadow plant species and soil microbial community composition, using phospholipid fatty acids (PLFAs) and high-throughput sequencing. We primarily focused on the root economics spectrum, as it provides a useful framework to examine plant strategies by integrating the co-ordination of belowground root traits along a resource acquisition-conservation trade-off axis. We found that the chemical axis of the fine root economics spectrum was positively related to fungal to bacterial ratios, but negatively to Gram-positive to Gram-negative bacterial ratios. However, this spectrum was unrelated to the relative abundance of functional guilds of soil fungi. Nevertheless, the relative abundance of arbuscular mycorrhizal fungi was positively correlated to root carbon content, but negatively to the numbers of root forks per root length. Our results suggest that the fine root economics spectrum is important for predicting broader groups of soil microorganisms (i.e. fungi and bacteria), while individual root traits may be more important for predicting soil microbial taxa and functional guilds.

Soil biotic and abiotic effects on seedling growth exhibit context-dependent interactions: evidence from a multi-country experiment on Pinus contorta invasion.

The success of invasive plants is influenced by many interacting factors, but evaluating multiple possible mechanisms of invasion success and elucidating the relative importance of abiotic and biotic drivers is challenging, and therefore rarely achieved. We used live, sterile or inoculated soil from different soil origins (native range and introduced range plantation; and invaded plots spanning three different countries) in a fully factorial design to simultaneously examine the influence of soil origin and soil abiotic and biotic factors on the growth of invasive Pinus contorta. Our results displayed significant context dependency in that certain soil abiotic conditions in the introduced ranges (soil nitrogen, phosphorus or carbon content) influenced responses to inoculation treatments. Our findings do not support the enemy release hypothesis or the enhanced mutualism hypothesis, as biota from native and plantation ranges promoted growth similarly. Instead, our results support the missed mutualism hypothesis, as biota from invasive ranges were the least beneficial for seedling growth. Our study provides a novel perspective on how variation in soil abiotic factors can influence plant-soil feedbacks for an invasive tree across broad biogeographical contexts.

Urbanization minimizes the effects of plant traits on soil provisioned ecosystem services across climatic regions.

An increasingly urbanized world is one of the most prominent examples of global environmental change. Across the globe, urban parks are designed and managed in a similar way, resulting in visually pleasing expansions of lawn interspersed with individually planted trees of varying appearances and functional traits. These large urban greenspaces have the capacity to provide various ecosystem services, including those associated with soil physicochemical properties. Our aim was to explore whether soil properties in urban parks diverge underneath vegetation producing labile or recalcitrant litter, and whether the impact is affected by climatic zone (from a boreal to temperate to tropical city). We also compared these properties to those in (semi)natural forests outside the cities to assess the influence of urbanization on plant-trait effects. We showed that vegetation type affected percentage soil organic matter (OM), total carbon (C) and total nitrogen (N), but inconsistently across climatic zones. Plant-trait effects were particularly weak in old parks in the boreal and temperate zones, whereas in young parks in these zones, soils underneath the two tree types accumulated significantly more OM, C and N compared to lawns. Within climatic zones, anthropogenic drivers dominated natural ones, with consistently lower values of organic-matter-related soil properties under trees producing labile or recalcitrant litter in parks compared to forests. The dominating effect of urbanization is also reflected in its ability to homogenize soil properties in parks across the three cities, especially in lawn soils and soils under trees irrespective of functional trait. Our study demonstrates that soil functions that relate to carbon and nitrogen dynamics-even in old urban greenspaces where plant-soil interactions have a long history-clearly diverged from those in natural ecosystems, implying a long-lasting influence of anthropogenic drivers on soil ecosystem services.

Rhizosphere control of soil nitrogen cycling: a key component of plant economic strategies.

Understanding how plant species influence soil nutrient cycling is a major theme in terrestrial ecosystem ecology. However, the prevailing paradigm has mostly focused on litter decomposition, while rhizosphere effects on soil organic matter (SOM) decomposition have attracted little attention. Using a dual 13 C/15 N labeling approach in a 'common garden' glasshouse experiment, we investigated how the economic strategies of 12 grassland plant species (graminoids, forbs and legumes) drive soil nitrogen (N) cycling via rhizosphere processes, and how this in turn affects plant N acquisition and growth. Acquisitive species with higher photosynthesis, carbon rhizodeposition and N uptake than conservative species induced a stronger acceleration of soil N cycling through rhizosphere priming of SOM decomposition. This allowed them to take up larger amounts of N and allocate it above ground to promote photosynthesis, thereby sustaining their faster growth. The N2 -fixation ability of legumes enhanced rhizosphere priming by promoting photosynthesis and rhizodeposition. Our study demonstrates that the economic strategies of plant species regulate a plant-soil carbon-nitrogen feedback operating through the rhizosphere. These findings provide novel mechanistic insights into how plant species with contrasting economic strategies sustain their nutrition and growth through regulating the cycling of nutrients by soil microbes in their rhizosphere.

Mosses modify effects of warmer and wetter conditions on tree seedlings at the alpine treeline.

Climate warming enables tree seedling establishment beyond the current alpine treeline, but to achieve this, seedlings have to establish within existing tundra vegetation. In tundra, mosses are a prominent feature, known to regulate soil temperature and moisture through their physical structure and associated water retention capacity. Moss presence and species identity might therefore modify the impact of increases in temperature and precipitation on tree seedling establishment at the arctic-alpine treeline. We followed Betula pubescens and Pinus sylvestris seedling survival and growth during three growing seasons in the field. Tree seedlings were transplanted along a natural precipitation gradient at the subarctic-alpine treeline in northern Sweden, into plots dominated by each of three common moss species and exposed to combinations of moss removal and experimental warming by open-top chambers (OTCs). Independent of climate, the presence of feather moss, but not Sphagnum, strongly supressed survival of both tree species. Positive effects of warming and precipitation on survival and growth of B. pubescens seedlings occurred in the absence of mosses and as expected, this was partly dependent on moss species. P. sylvestris survival was greatest at high precipitation, and this effect was more pronounced in Sphagnum than in feather moss plots irrespective of whether the mosses had been removed or not. Moss presence did not reduce the effects of OTCs on soil temperature. Mosses therefore modified seedling response to climate through other mechanisms, such as altered competition or nutrient availability. We conclude that both moss presence and species identity pose a strong control on seedling establishment at the alpine treeline, and that in some cases mosses weaken climate-change effects on seedling establishment. Changes in moss abundance and species composition therefore have the potential to hamper treeline expansion induced by climate warming.

Effects of plant functional group removal on structure and function of soil communities across contrasting ecosystems.

Loss of plant diversity has an impact on ecosystems worldwide, but we lack a mechanistic understanding of how this loss may influence below-ground biota and ecosystem functions across contrasting ecosystems in the long term. We used the longest running biodiversity manipulation experiment across contrasting ecosystems in existence to explore the below-ground consequences of 19 years of plant functional group removals for each of 30 contrasting forested lake islands in northern Sweden. We found that, against expectations, the effects of plant removals on the communities of key groups of soil organisms (bacteria, fungi and nematodes), and organic matter quality and soil ecosystem functioning (decomposition and microbial activity) were relatively similar among islands that varied greatly in productivity and soil fertility. This highlights that, in contrast to what has been shown for plant productivity, plant biodiversity loss effects on below-ground functions can be relatively insensitive to environmental context or variation among widely contrasting ecosystems.

Comparison of plant-soil feedback experimental approaches for testing soil biotic interactions among ecosystems.

The study of interactions and feedbacks between plants and soils is a rapidly expanding research area, and a primary tool used in this field is to perform glasshouse experiments where soil biota are manipulated. Recently, there has been vigorous debate regarding the correctness of methods for carrying out these types of experiment, and specifically whether it is legitimate to mix soils from different sites or plots (mixed soil sampling, MSS) or not (independent soil sampling, ISS) to create either soil inoculum treatments or subjects. We performed the first empirical comparison of MSS vs ISS approaches by comparing growth of two boreal tree species (Picea abies and Pinus sylvestris) in soils originating from 10 sites near the boreal forest limit in northern Sweden, and 10 sites in the subarctic region where boreal forests may potentially expand as a result of climate change. We found no consistent differences in the conclusions that we reached whether we used MSS or ISS approaches. We propose that researchers should not choose a soil handling method based on arguments that one method is inherently more correct than the other, but rather that method choice should be based on correct alignment with specific research questions and goals.

Biodiversity loss and its impact on humanity.

The most unique feature of Earth is the existence of life, and the most extraordinary feature of life is its diversity. Approximately 9 million types of plants, animals, protists and fungi inhabit the Earth. So, too, do 7 billion people. Two decades ago, at the first Earth Summit, the vast majority of the world's nations declared that human actions were dismantling the Earth's ecosystems, eliminating genes, species and biological traits at an alarming rate. This observation led to the question of how such loss of biological diversity will alter the functioning of ecosystems and their ability to provide society with the goods and services needed to prosper.