Anthropogenic and environmental factors partly co-determine the level, composition and temporal variation of beach debris.

The accumulation of human-derived waste on our coasts is an escalating phenomenon, yet the relative importance and potential interactions among its main drivers are not fully understood. We used citizen-science standardized collections to investigate how anthropogenic and environmental factors influence the level, composition, and temporal variation of beach debris. An average of 58 kg and 803 items/100 m, dominated by single-use items of land (rather than sea) origin, were collected in the 881 beaches sampled. Interactions between anthropogenic and environmental factors (e.g., human use × beach substrate) were the strongest predictors of beach debris, accounting for 34% of the variance explained in its amount and composition. Beach debris showed a highly stochastic temporal variation (adjusted R2 = 0.05), partly determined by interactions between different local and landscape anthropogenic pressures. Our results show that both environmental and anthropogenic factors (at the local and landscape scale) co-determine the level and composition of beach debris. We emphasize the potential of citizen-science to inform environmental policy, showing that land-originated single-use items dominate beach debris, and the importance of considering their multiple anthropogenic and environmental drivers to improve our low predictive power regarding their spatio-temporal distribution.

Nitrogen availability and plant functional composition modify biodiversity-multifunctionality relationships.

Biodiversity typically increases multiple ecosystem functions simultaneously (multifunctionality) but variation in the strength and direction of biodiversity effects between studies suggests context dependency. To determine how different factors modulate the diversity effect on multifunctionality, we established a large grassland experiment manipulating plant species richness, resource addition, functional composition (exploitative vs. conservative species), functional diversity and enemy abundance. We measured ten above- and belowground functions and calculated ecosystem multifunctionality. Species richness and functional diversity both increased multifunctionality, but their effects were context dependent. Richness increased multifunctionality when communities were assembled with fast-growing species. This was because slow species were more redundant in their functional effects, whereas different fast species promoted different functions. Functional diversity also increased multifunctionality but this effect was dampened by nitrogen enrichment and enemy presence. Our study suggests that a shift towards fast-growing communities will not only alter ecosystem functioning but also the strength of biodiversity-functioning relationships.

Rewilding soil-disturbing vertebrates to rehabilitate degraded landscapes: benefits and risks.

Soil-disturbing animals are common globally and play important roles in creating and maintaining healthy functional soils and landscapes. Yet many of these animals are threatened or locally extinct due to habitat loss, predation by non-native animals or poaching and poisoning. Some reintroduction and rewilding programmes have as their core aims to increase animal populations and reinstate processes that have been lost due to their extirpation. Here we use a meta-analytical approach to review the effects of soil-disturbing vertebrates on ecosystem processes, and advance the argument that they can be used to rehabilitate degraded ecosystems by altering mainly composition and function, but with fewer positive effects on structure. We describe four examples where the loss or reintroduction of soil-disturbing vertebrates leads to ecosystem state changes and highlight the role of spatial scale, covarying management changes, and species co-occurrence in modulating their effects. We discuss the advantages and disadvantages of using soil-disturbing vertebrates over mechanized engineering approaches such as pitting and furrowing, considering some advantages to include more self-sustainable and heterogeneous disturbances, creation of new habitats and added recreational values. Finally, we identify key knowledge gaps in our understanding of the use of soil-disturbing vertebrates for rehabilitating degraded ecosystems.

Thinning and plantation of resprouting species redirect overstocked pine stands towards more functional communities in the Mediterranean basin.

Post-fire regeneration in Pinus halepensis' forests, one of the most abundant vegetation types in the Mediterranean basin, often generates overstocked and vulnerable stands. They accumulate a high fuel load, increasing the risk of further fires, and present high levels of vulnerability due to their reduced seed production. In addition, these dense stands substantially reduce the availability of light and nutrients, which may hinder the recruitment of other species, often generating mono-specific and homogeneous stands, which potentially supply fewer ecosystem services than mixed forests with more heterogeneous structures. In these dense pine stands, management is of high priority to reduce fire hazards and promote their functionality. In overstocked pine stands (>75,000 trees·ha-1), we assessed the long-term effects (10 years) of two thinning levels (600 and 1200 trees·ha-1), in combination with the plantation of Quercus faginea (a resprouter species typical of advanced successional stages in our study area) on 28 above and belowground ecosystem attributes, including fire hazard. After ten years, thinning and plantation interacted to enhance ecosystem attributes associated with disturbance regulation and biodiversity conservation (up to 200%) and food production (up to 90%), while no effects were observed on those attributes related to carbon sequestration and supporting services. These effects were mainly driven by aboveground attributes, as they responded more strongly to our treatments than those belowground. Our results are relevant for the restoration of Mediterranean degraded ecosystems, and show that tree thinning in overstocked pine stands, combined with the plantation of resprouter species, may not only reduce fire risks and accelerate post-fire succession but also enhance the supply of multiple ecosystem services in the long run.

Biogeography of global drylands.

Despite their extent and socio-ecological importance, a comprehensive biogeographical synthesis of drylands is lacking. Here we synthesize the biogeography of key organisms (vascular and nonvascular vegetation and soil microorganisms), attributes (functional traits, spatial patterns, plant-plant and plant-soil interactions) and processes (productivity and land cover) across global drylands. These areas have a long evolutionary history, are centers of diversification for many plant lineages and include important plant diversity hotspots. This diversity captures a strikingly high portion of the variation in leaf functional diversity observed globally. Part of this functional diversity is associated with the large variation in response and effect traits in the shrubs encroaching dryland grasslands. Aridity and its interplay with the traits of interacting plant species largely shape biogeographical patterns in plant-plant and plant-soil interactions, and in plant spatial patterns. Aridity also drives the composition of biocrust communities and vegetation productivity, which shows large geographical variation. We finish our review by discussing major research gaps, which include: studying regular vegetation spatial patterns; establishing large-scale plant and biocrust field surveys assessing individual-level trait measurements; knowing whether the impacts of plant-plant and plant-soil interactions on biodiversity are predictable; and assessing how elevated CO2 modulates future aridity conditions and plant productivity.

Land-use intensity alters networks between biodiversity, ecosystem functions, and services.

Land-use intensification can increase provisioning ecosystem services, such as food and timber production, but it also drives changes in ecosystem functioning and biodiversity loss, which may ultimately compromise human wellbeing. To understand how changes in land-use intensity affect the relationships between biodiversity, ecosystem functions, and services, we built networks from correlations between the species richness of 16 trophic groups, 10 ecosystem functions, and 15 ecosystem services. We evaluated how the properties of these networks varied across land-use intensity gradients for 150 forests and 150 grasslands. Land-use intensity significantly affected network structure in both habitats. Changes in connectance were larger in forests, while changes in modularity and evenness were more evident in grasslands. Our results show that increasing land-use intensity leads to more homogeneous networks with less integration within modules in both habitats, driven by the belowground compartment in grasslands, while forest responses to land management were more complex. Land-use intensity strongly altered hub identity and module composition in both habitats, showing that the positive correlations of provisioning services with biodiversity and ecosystem functions found at low land-use intensity levels, decline at higher intensity levels. Our approach provides a comprehensive view of the relationships between multiple components of biodiversity, ecosystem functions, and ecosystem services and how they respond to land use. This can be used to identify overall changes in the ecosystem, to derive mechanistic hypotheses, and it can be readily applied to further global change drivers.

Global ecosystem thresholds driven by aridity.

Aridity, which is increasing worldwide because of climate change, affects the structure and functioning of dryland ecosystems. Whether aridification leads to gradual (versus abrupt) and systemic (versus specific) ecosystem changes is largely unknown. We investigated how 20 structural and functional ecosystem attributes respond to aridity in global drylands. Aridification led to systemic and abrupt changes in multiple ecosystem attributes. These changes occurred sequentially in three phases characterized by abrupt decays in plant productivity, soil fertility, and plant cover and richness at aridity values of 0.54, 0.7, and 0.8, respectively. More than 20% of the terrestrial surface will cross one or several of these thresholds by 2100, which calls for immediate actions to minimize the negative impacts of aridification on essential ecosystem services for the more than 2 billion people living in drylands.

Phylogenetic, functional, and taxonomic richness have both positive and negative effects on ecosystem multifunctionality.

Biodiversity encompasses multiple attributes such as the richness and abundance of species (taxonomic diversity), the presence of different evolutionary lineages (phylogenetic diversity), and the variety of growth forms and resource use strategies (functional diversity). These biodiversity attributes do not necessarily relate to each other and may have contrasting effects on ecosystem functioning. However, how they simultaneously influence the provision of multiple ecosystem functions related to carbon, nitrogen, and phosphorus cycling (multifunctionality) remains unknown. We evaluated the effects of the taxonomic, phylogenetic, and functional attributes of dominant (mass ratio effects) and subordinate (richness effect) plant species on the multifunctionality of 123 drylands from six continents. Our results highlight the importance of the phylogenetic and functional attributes of subordinate species as key drivers of multifunctionality. In addition to a higher taxonomic richness, we found that simultaneously increasing the richness of early diverging lineages and the functional redundancy between species increased multifunctionality. In contrast, the richness of most recent evolutionary lineages and the functional and phylogenetic attributes of dominant plant species (mass ratio effects) were weakly correlated with multifunctionality. However, they were important drivers of individual nutrient cycles. By identifying which biodiversity attributes contribute the most to multifunctionality, our results can guide restoration efforts aiming to maximize either multifunctionality or particular nutrient cycles, a critical step to combat dryland desertification worldwide.

Multifunctionality debt in global drylands linked to past biome and climate.

Past vegetation and climatic conditions are known to influence current biodiversity patterns. However, whether their legacy effects affect the provision of multiple ecosystem functions, that is, multifunctionality, remains largely unknown. Here we analyzed soil nutrient stocks and their transformation rates in 236 drylands from six continents to evaluate the associations between current levels of multifunctionality and legacy effects of the Last Glacial Maximum (LGM) desert biome distribution and climate. We found that past desert distribution and temperature legacy, defined as increasing temperature from LGM, were negatively correlated with contemporary multifunctionality even after accounting for predictors such as current climate, soil texture, plant species richness, and site topography. Ecosystems that have been deserts since the LGM had up to 30% lower contemporary multifunctionality compared with those that were nondeserts during the LGM. In addition, ecosystems that experienced higher warming rates since the LGM had lower contemporary multifunctionality than those suffering lower warming rates, with a ~9% reduction per extra degree Celsius. Past desert distribution and temperature legacies had direct negative effects, while temperature legacy also had indirect (via soil sand content) negative effects on multifunctionality. Our results indicate that past biome and climatic conditions have left a strong "functionality debt" in global drylands. They also suggest that ongoing warming and expansion of desert areas may leave a strong fingerprint in the future functioning of dryland ecosystems worldwide that needs to be considered when establishing management actions aiming to combat land degradation and desertification.

Specialisation and diversity of multiple trophic groups are promoted by different forest features.

While forest management strongly influences biodiversity, it remains unclear how the structural and compositional changes caused by management affect different community dimensions (e.g. richness, specialisation, abundance or completeness) and how this differs between taxa. We assessed the effects of nine forest features (representing stand structure, heterogeneity and tree composition) on thirteen above- and belowground trophic groups of plants, animals, fungi and bacteria in 150 temperate forest plots differing in their management type. Canopy cover decreased light resources, which increased community specialisation but reduced overall diversity and abundance. Features increasing resource types and diversifying microhabitats (admixing of oaks and conifers) were important and mostly affected richness. Belowground groups responded differently to those aboveground and had weaker responses to most forest features. Our results show that we need to consider forest features rather than broad management types and highlight the importance of considering several groups and community dimensions to better inform conservation.

Multiple forest attributes underpin the supply of multiple ecosystem services.

Trade-offs and synergies in the supply of forest ecosystem services are common but the drivers of these relationships are poorly understood. To guide management that seeks to promote multiple services, we investigated the relationships between 12 stand-level forest attributes, including structure, composition, heterogeneity and plant diversity, plus 4 environmental factors, and proxies for 14 ecosystem services in 150 temperate forest plots. Our results show that forest attributes are the best predictors of most ecosystem services and are also good predictors of several synergies and trade-offs between services. Environmental factors also play an important role, mostly in combination with forest attributes. Our study suggests that managing forests to increase structural heterogeneity, maintain large trees, and canopy gaps would promote the supply of multiple ecosystem services. These results highlight the potential for forest management to encourage multifunctional forests and suggest that a coordinated landscape-scale strategy could help to mitigate trade-offs in human-dominated landscapes.

Publisher Correction: Redefining ecosystem multifunctionality.

In the version of this Perspective originally published, in the figure in Box 3 the middle panel of the top row was incorrectly labelled '50% threshold-plus'; it should have read '50% threshold'. This has now been corrected.

Redefining ecosystem multifunctionality.

Recent years have seen a surge of interest in ecosystem multifunctionality, a concept that has developed in the largely separate fields of biodiversity-ecosystem function and land management research. Here we discuss the merit of the multifunctionality concept, the advances it has delivered, the challenges it faces and solutions to these challenges. This involves the redefinition of multifunctionality as a property that exists at two levels: ecosystem function multifunctionality and ecosystem service multifunctionality. The framework presented provides a road map for the development of multifunctionality measures that are robust, quantifiable and relevant to both fundamental ecological science and ecosystem management.

Author Correction: Plant spatial patterns identify alternative ecosystem multifunctionality states in global drylands.

In the version of this Article originally published, the values of two of the functions used to calculate the multifunctionality index were incorrect, which affected Figs 3,4 of the main article and Supplementary Figs 3,4,5,6,9. Please see the correction notice for full details.

Plant spatial patterns identify alternative ecosystem multifunctionality states in global drylands.

The response of drylands to environmental gradients can be abrupt rather than gradual. These shifts largely occur unannounced and are difficult to reverse once they happen; their prompt detection is of crucial importance. The distribution of vegetation patch sizes may indicate the proximity to these shifts, but the use of this metric is hampered by a lack of large-scale studies relating these distributions to the provision of multiple ecosystem functions (multifunctionality) and comparing them to other ecosystem attributes, such as total plant cover. Here we sampled 115 dryland ecosystems across the globe and related their vegetation attributes (cover and patch size distributions) to multifunctionality. Multifunctionality followed a bimodal distribution across our sites, suggesting alternative states in the functioning of drylands. Although plant cover was the strongest predictor of multifunctionality when linear analyses were used, only patch size distributions reflected the bimodal distribution of multifunctionality observed. Differences in the coupling between nutrient cycles and in the importance of self-organizing biotic processes characterized the two multifunctionality states observed. Our findings support the use of vegetation patterns as indicators of ecosystem functioning in drylands and pave the way for developing effective strategies to monitor desertification processes.

Ecosystem structure, function, and composition in rangelands are negatively affected by livestock grazing.

Reports of positive or neutral effects of grazing on plant species richness have prompted calls for livestock grazing to be used as a tool for managing land for conservation. Grazing effects, however, are likely to vary among different response variables, types, and intensity of grazing, and across abiotic conditions. We aimed to examine how grazing affects ecosystem structure, function, and composition. We compiled a database of 7615 records reporting an effect of grazing by sheep and cattle on 278 biotic and abiotic response variables for published studies across Australia. Using these data, we derived three ecosystem measures based on structure, function, and composition, which were compared against six contrasts of grazing pressure, ranging from low to heavy, two different herbivores (sheep, cattle), and across three different climatic zones. Grazing reduced structure (by 35%), function (24%), and composition (10%). Structure and function (but not composition) declined more when grazed by sheep and cattle together than sheep alone. Grazing reduced plant biomass (40%), animal richness (15%), and plant and animal abundance, and plant and litter cover (25%), but had no effect on plant richness nor soil function. The negative effects of grazing on plant biomass, plant cover, and soil function were more pronounced in drier environments. Grazing effects on plant and animal richness and composition were constant, or even declined, with increasing aridity. Our study represents a comprehensive continental assessment of the implications of grazing for managing Australian rangelands. Grazing effects were largely negative, even at very low levels of grazing. Overall, our results suggest that livestock grazing in Australia is unlikely to produce positive outcomes for ecosystem structure, function, and composition or even as a blanket conservation tool unless reduction in specific response variables is an explicit management objective.

Biodiversity at multiple trophic levels is needed for ecosystem multifunctionality.

Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provide the multiple services on which humans depend. However, experiments necessarily simplify the complexity of natural ecosystems and will normally control for other important drivers of ecosystem functioning, such as the environment or land use. In addition, existing studies typically focus on the diversity of single trophic groups, neglecting the fact that biodiversity loss occurs across many taxa and that the functional effects of any trophic group may depend on the abundance and diversity of others. Here we report analysis of the relationships between the species richness and abundance of nine trophic groups, including 4,600 above- and below-ground taxa, and 14 ecosystem services and functions and with their simultaneous provision (or multifunctionality) in 150 grasslands. We show that high species richness in multiple trophic groups (multitrophic richness) had stronger positive effects on ecosystem services than richness in any individual trophic group; this includes plant species richness, the most widely used measure of biodiversity. On average, three trophic groups influenced each ecosystem service, with each trophic group influencing at least one service. Multitrophic richness was particularly beneficial for 'regulating' and 'cultural' services, and for multifunctionality, whereas a change in the total abundance of species or biomass in multiple trophic groups (the multitrophic abundance) positively affected supporting services. Multitrophic richness and abundance drove ecosystem functioning as strongly as abiotic conditions and land-use intensity, extending previous experimental results to real-world ecosystems. Primary producers, herbivorous insects and microbial decomposers seem to be particularly important drivers of ecosystem functioning, as shown by the strong and frequent positive associations of their richness or abundance with multiple ecosystem services. Our results show that multitrophic richness and abundance support ecosystem functioning, and demonstrate that a focus on single groups has led to researchers to greatly underestimate the functional importance of biodiversity.

Environmental correlates of species rank - abundance distributions in global drylands.

Theoretical models predict lognormal species abundance distributions (SADs) in stable and productive environments, with log-series SADs in less stable, dispersal driven communities. We studied patterns of relative species abundances of perennial vascular plants in global dryland communities to: i) assess the influence of climatic and soil characteristics on the observed SADs, ii) infer how environmental variability influences relative abundances, and iii) evaluate how colonisation dynamics and environmental filters shape abundance distributions. We fitted lognormal and log-series SADs to 91 sites containing at least 15 species of perennial vascular plants. The dependence of species relative abundances on soil and climate variables was assessed using general linear models. Irrespective of habitat type and latitude, the majority of the SADs (70.3%) were best described by a lognormal distribution. Lognormal SADs were associated with low annual precipitation, higher aridity, high soil carbon content, and higher variability of climate variables and soil nitrate. Our results do not corroborate models predicting the prevalence of log-series SADs in dryland communities. As lognormal SADs were particularly associated with sites with drier conditions and a higher environmental variability, we reject models linking lognormality to environmental stability and high productivity conditions. Instead our results point to the prevalence of lognormal SADs in heterogeneous environments, allowing for more evenly distributed plant communities, or in stressful ecosystems, which are generally shaped by strong habitat filters and limited colonisation. This suggests that drylands may be resilient to environmental changes because the many species with intermediate relative abundances could take over ecosystem functioning if the environment becomes suboptimal for dominant species.

Locally rare species influence grassland ecosystem multifunctionality.

Species diversity promotes the delivery of multiple ecosystem functions (multifunctionality). However, the relative functional importance of rare and common species in driving the biodiversity-multifunctionality relationship remains unknown. We studied the relationship between the diversity of rare and common species (according to their local abundances and across nine different trophic groups), and multifunctionality indices derived from 14 ecosystem functions on 150 grasslands across a land-use intensity (LUI) gradient. The diversity of above- and below-ground rare species had opposite effects, with rare above-ground species being associated with high levels of multifunctionality, probably because their effects on different functions did not trade off against each other. Conversely, common species were only related to average, not high, levels of multifunctionality, and their functional effects declined with LUI. Apart from the community-level effects of diversity, we found significant positive associations between the abundance of individual species and multifunctionality in 6% of the species tested. Species-specific functional effects were best predicted by their response to LUI: species that declined in abundance with land use intensification were those associated with higher levels of multifunctionality. Our results highlight the importance of rare species for ecosystem multifunctionality and help guiding future conservation priorities.