Divergent ecological responses to typhoon disturbance revealed via landscape-scale acoustic monitoring.

Climate change is increasing the frequency, intensity, and duration of extreme weather events across the globe. Understanding the capacity for ecological communities to withstand and recover from such events is critical. Typhoons are extreme weather events that are expected to broadly homogenize ecological dynamics through structural damage to vegetation and longer-term effects of salinization. Given their unpredictable nature, monitoring ecological responses to typhoons is challenging, particularly for mobile animals such as birds. Here, we report spatially variable ecological responses to typhoons across terrestrial landscapes. Using a high temporal resolution passive acoustic monitoring network across 24 sites on the subtropical island of Okinawa, Japan, we found that typhoons elicit divergent ecological responses among Okinawa's diverse terrestrial habitats, as indicated by increased spatial variability of biological sound production (biophony) and individual species detections. This suggests that soniferous communities are capable of a diversity of different responses to typhoons. That is, spatial insurance effects among local ecological communities provide resilience to typhoons at the landscape scale. Even though site-level typhoon impacts on soundscapes and bird detections were not particularly strong, monitoring at scale with high temporal resolution across a broad spatial extent nevertheless enabled detection of spatial heterogeneity in typhoon responses. Further, species-level responses mirrored those of acoustic indices, underscoring the utility of such indices for revealing insight into fundamental questions concerning disturbance and stability. Our findings demonstrate the significant potential of landscape-scale acoustic sensor networks to capture the understudied ecological impacts of unpredictable extreme weather events.

Breakdown in seasonal dynamics of subtropical ant communities with land-cover change.

Concerns about widespread human-induced declines in insect populations are mounting, yet little is known about how land-use change modifies both the trends and variability of insect communities, particularly in understudied regions. Here, we examine how the seasonal activity patterns of ants-key drivers of terrestrial ecosystem functioning-vary with anthropogenic land-cover change on a subtropical island landscape, and whether differences in temperature or species composition can explain observed patterns. Using trap captures sampled biweekly over 2 years from a biodiversity monitoring network covering Okinawa Island, Japan, we processed 1.2 million individuals and reconstructed activity patterns within and across habitat types. Forest communities exhibited greater temporal variability of activity than those in more developed areas. Using time-series decomposition to deconstruct this pattern, we found that sites with greater human development exhibited ant communities with diminished seasonality, reduced synchrony and higher stochasticity compared with sites with greater forest cover. Our results cannot be explained by variation in regional or site temperature patterns, or by differences in species richness or composition among sites. Our study raises the possibility that disruptions to natural seasonal patterns of functionally key insect communities may comprise an important and underappreciated consequence of global environmental change that must be better understood across Earth's biomes.

Multidimensional responses of grassland stability to eutrophication.

Eutrophication usually impacts grassland biodiversity, community composition, and biomass production, but its impact on the stability of these community aspects is unclear. One challenge is that stability has many facets that can be tightly correlated (low dimensionality) or highly disparate (high dimensionality). Using standardized experiments in 55 grassland sites from a globally distributed experiment (NutNet), we quantify the effects of nutrient addition on five facets of stability (temporal invariability, resistance during dry and wet growing seasons, recovery after dry and wet growing seasons), measured on three community aspects (aboveground biomass, community composition, and species richness). Nutrient addition reduces the temporal invariability and resistance of species richness and community composition during dry and wet growing seasons, but does not affect those of biomass. Different stability measures are largely uncorrelated under both ambient and eutrophic conditions, indicating consistently high dimensionality. Harnessing the dimensionality of ecological stability provides insights for predicting grassland responses to global environmental change.

NEAL: an open-source tool for audio annotation.

Passive acoustic monitoring is used widely in ecology, biodiversity, and conservation studies. Data sets collected via acoustic monitoring are often extremely large and built to be processed automatically using artificial intelligence and machine learning models, which aim to replicate the work of domain experts. These models, being supervised learning algorithms, need to be trained on high quality annotations produced by experts. Since the experts are often resource-limited, a cost-effective process for annotating audio is needed to get maximal use out of the data. We present an open-source interactive audio data annotation tool, NEAL (Nature+Energy Audio Labeller). Built using R and the associated Shiny framework, the tool provides a reactive environment where users can quickly annotate audio files and adjust settings that automatically change the corresponding elements of the user interface. The app has been designed with the goal of having both expert birders and citizen scientists contribute to acoustic annotation projects. The popularity and flexibility of R programming in bioacoustics means that the Shiny app can be modified for other bird labelling data sets, or even to generic audio labelling tasks. We demonstrate the app by labelling data collected from wind farm sites across Ireland.

Environmental heterogeneity modulates the effect of plant diversity on the spatial variability of grassland biomass.

Plant productivity varies due to environmental heterogeneity, and theory suggests that plant diversity can reduce this variation. While there is strong evidence of diversity effects on temporal variability of productivity, whether this mechanism extends to variability across space remains elusive. Here we determine the relationship between plant diversity and spatial variability of productivity in 83 grasslands, and quantify the effect of experimentally increased spatial heterogeneity in environmental conditions on this relationship. We found that communities with higher plant species richness (alpha and gamma diversity) have lower spatial variability of productivity as reduced abundance of some species can be compensated for by increased abundance of other species. In contrast, high species dissimilarity among local communities (beta diversity) is positively associated with spatial variability of productivity, suggesting that changes in species composition can scale up to affect productivity. Experimentally increased spatial environmental heterogeneity weakens the effect of plant alpha and gamma diversity, and reveals that beta diversity can simultaneously decrease and increase spatial variability of productivity. Our findings unveil the generality of the diversity-stability theory across space, and suggest that reduced local diversity and biotic homogenization can affect the spatial reliability of key ecosystem functions.

Reconciling climate action with the need for biodiversity protection, restoration and rehabilitation.

Globally, we are faced with a climate crisis that requires urgent transition to a low-carbon economy. Simultaneously, the biodiversity crisis demands equally urgent action to prevent further species loss and promote restoration and rehabilitation of ecosystems. Climate action itself must prevent further pressures on biodiversity and options for synergistic gains for both climate and biodiversity change mitigation and adaptation need to be explored and implemented. Here, we review the key potential impacts of climate mitigation measures in energy and land-use on biodiversity, including the development of renewable energy such as offshore and onshore wind, solar, and bioenergy. We also assess the potential impacts of climate action driven afforestation and native habitat rehabilitation and restoration. We apply our findings to Ireland as a unique case-study as the government develops a coordinated response to climate and biodiversity change through declaration of a joint climate and biodiversity emergency and inclusion of biodiversity in key climate change legislation and the national Climate Action Plan. However, acknowledgement of these intertwined crises is only a first step; implementation of synergistic solutions requires careful planning. We demonstrate how synergy between climate and biodiversity action can be gained through explicit consideration of the effects of climate change mitigation strategies, such as energy infrastructure development and land-use change, on biodiversity. We identify several potential "win-win" strategies for both climate mitigation and biodiversity conservation. For Ireland, these include increasing offshore wind capacity, rehabilitating natural areas surrounding onshore wind turbines, and limiting the development of solar photovoltaics to the built environment. Ultimately, climate mitigation should be implemented in a "Right Action, Right Place" framework to maximise positive biodiversity benefits. This review provides one of the first examples of how national climate actions can be implemented in a biodiversity-conscious way to initiate discussion about synergistic solutions for both climate and biodiversity.

A function-based typology for Earth's ecosystems.

As the United Nations develops a post-2020 global biodiversity framework for the Convention on Biological Diversity, attention is focusing on how new goals and targets for ecosystem conservation might serve its vision of 'living in harmony with nature'1,2. Advancing dual imperatives to conserve biodiversity and sustain ecosystem services requires reliable and resilient generalizations and predictions about ecosystem responses to environmental change and management3. Ecosystems vary in their biota4, service provision5 and relative exposure to risks6, yet there is no globally consistent classification of ecosystems that reflects functional responses to change and management. This hampers progress on developing conservation targets and sustainability goals. Here we present the International Union for Conservation of Nature (IUCN) Global Ecosystem Typology, a conceptually robust, scalable, spatially explicit approach for generalizations and predictions about functions, biota, risks and management remedies across the entire biosphere. The outcome of a major cross-disciplinary collaboration, this novel framework places all of Earth's ecosystems into a unifying theoretical context to guide the transformation of ecosystem policy and management from global to local scales. This new information infrastructure will support knowledge transfer for ecosystem-specific management and restoration, globally standardized ecosystem risk assessments, natural capital accounting and progress on the post-2020 global biodiversity framework.

Linking changes in species composition and biomass in a globally distributed grassland experiment.

Global change drivers, such as anthropogenic nutrient inputs, are increasing globally. Nutrient deposition simultaneously alters plant biodiversity, species composition and ecosystem processes like aboveground biomass production. These changes are underpinned by species extinction, colonisation and shifting relative abundance. Here, we use the Price equation to quantify and link the contributions of species that are lost, gained or that persist to change in aboveground biomass in 59 experimental grassland sites. Under ambient (control) conditions, compositional and biomass turnover was high, and losses (i.e. local extinctions) were balanced by gains (i.e. colonisation). Under fertilisation, the decline in species richness resulted from increased species loss and decreases in species gained. Biomass increase under fertilisation resulted mostly from species that persist and to a lesser extent from species gained. Drivers of ecological change can interact relatively independently with diversity, composition and ecosystem processes and functions such as aboveground biomass due to the individual contributions of species lost, gained or persisting.

Evolutionary history of grazing and resources determine herbivore exclusion effects on plant diversity.

Ecological models predict that the effects of mammalian herbivore exclusion on plant diversity depend on resource availability and plant exposure to ungulate grazing over evolutionary time. Using an experiment replicated in 57 grasslands on six continents, with contrasting evolutionary history of grazing, we tested how resources (mean annual precipitation and soil nutrients) determine herbivore exclusion effects on plant diversity, richness and evenness. Here we show that at sites with a long history of ungulate grazing, herbivore exclusion reduced plant diversity by reducing both richness and evenness and the responses of richness and diversity to herbivore exclusion decreased with mean annual precipitation. At sites with a short history of grazing, the effects of herbivore exclusion were not related to precipitation but differed for native and exotic plant richness. Thus, plant species' evolutionary history of grazing continues to shape the response of the world's grasslands to changing mammalian herbivory.

Alternate patterns of temperature variation bring about very different disease outcomes at different mean temperatures.

The dynamics of host-parasite interactions are highly temperature-dependent and may be modified by increasing frequency and intensity of climate-driven heat events. Here, we show that altered patterns of temperature variance lead to an almost order-of-magnitude shift in thermal performance of host and pathogen life-history traits over and above the effects of mean temperature and, moreover, that different temperature regimes affect these traits differently. We found that diurnal fluctuations of ±3°C lowered infection rates and reduced spore burden compared to constant temperatures in our focal host Daphnia magna exposed to the microsporidium parasite Ordospora colligata. In contrast, a 3-day heatwave (+6°C) did not affect infection rates, but increased spore burden (relative to constant temperatures with the same mean) at 16°C, while reducing burden at higher temperatures. We conclude that changing patterns of climate variation, superimposed on shifts in mean temperatures due to global warming, may have profound and unanticipated effects on disease dynamics.

Global warming is increasing average temperatures and causing extreme temperature fluctuations and heatwaves. These changes may affect when, where, and how often infectious disease outbreaks occur. This could have profound impacts on agriculture, human health, and wildlife. Studying how extreme temperatures or temperature fluctuations alter infections in laboratory animals may help scientists to better understand the impact of climate change on disease. A small aquatic invertebrate, such as a water flea, is one good candidate for such studies. These tiny creatures can be grown in small glass jars in temperature-controlled aquariums. Kunze, Luijckx et al. show that temperature fluctuations and heat waves have complex effects on parasitic infections in water fleas. In the experiments, water fleas housed with a parasite that infects them were exposed to constant temperatures, fluctuating temperatures, or three-day heatwaves, while being kept at a broad range of mean water temperatures. Then, Kunze, Luijckx et al. measured how these conditions affected the water fleas' longevity, reproduction, and parasite infections. This revealed that temperature variations had a unique effect on the life span, and reproduction and infection rates of the water fleas, depending on the average water temperature the animals were kept at. Heatwaves drastically increased the number of parasites in the water fleas at an average water temperature of 16 °C but had no effect at all or decreased the number of parasites at 19 °C and 22 °C, respectively. Similarly, at high average water temperatures (>24 °C), temperature fluctuations reduced the number of water fleas infected with parasites and the number of parasites in each infected flea. Moreover, the maximum temperature at which parasites were able to cause infections was 5 °C lower under fluctuating temperatures than under constant temperatures. Kunze and Luijckx et al. show that consistent high temperatures, temperature changes, extreme weather events, and mean water temperature affect disease outcomes in water fleas. More studies are needed to assess how temperature variations change the course of diseases in other organisms and to understand the underlying mechanisms. Learning more about disease-temperature interactions will help scientists predict climate change-driven disease outbreaks.

Nutrients and herbivores impact grassland stability across spatial scales through different pathways.

Nutrients and herbivores are well-known drivers of grassland diversity and stability in local communities. However, whether they interact to impact the stability of aboveground biomass and whether these effects depend on spatial scales remain unknown. It is also unclear whether nutrients and herbivores impact stability via different facets of plant diversity including species richness, evenness, and changes in community composition through time and space. We used a replicated experiment adding nutrients and excluding herbivores for 5 years in 34 global grasslands to explore these questions. We found that both nutrient addition and herbivore exclusion alone reduced stability at the larger spatial scale (aggregated local communities; gamma stability), but through different pathways. Nutrient addition reduced gamma stability primarily by increasing changes in local community composition over time, which was mainly driven by species replacement. Herbivore exclusion reduced gamma stability primarily by decreasing asynchronous dynamics among local communities (spatial asynchrony). Their interaction weakly increased gamma stability by increasing spatial asynchrony. Our findings indicate that disentangling the processes operating at different spatial scales may improve conservation and management aiming at maintaining the ability of ecosystems to reliably provide functions and services for humanity.

Predators mitigate the destabilising effects of heatwaves on multitrophic stream communities.

Amidst the global extinction crisis, climate change will expose ecosystems to more frequent and intense extreme climatic events, such as heatwaves. Yet, whether predator species loss-a prevailing characteristic of the extinction crisis-will exacerbate the ecological consequences of extreme climatic events remains largely unknown. Here, we show that the loss of predator species can interact with heatwaves to moderate the compositional stability of ecosystems. We exposed multitrophic stream communities, with and without a dominant predator species, to realistic current and future heatwaves and found that heatwaves destabilised algal communities by homogenising them in space. However, this happened only when the predator was absent. Additional heatwave impacts on multiple aspects of stream communities, including changes to the structure of algal and macroinvertebrate communities, as well as total algal biomass and its temporal variability, were not apparent during heatwaves and emerged only after the heatwaves had passed. Taken together, our results suggest that the ecological consequences of heatwaves can amplify over time as their impacts propagate through biological interaction networks, but the presence of predators can help to buffer such impacts. These findings underscore the importance of conserving trophic structure, and highlight the potential for species extinctions to amplify the effects of climate change and extreme events.

Soil properties as key predictors of global grassland production: Have we overlooked micronutrients?

Fertilisation experiments have demonstrated that nutrient availability is a key determinant of biomass production and carbon sequestration in grasslands. However, the influence of nutrients in explaining spatial variation in grassland biomass production has rarely been assessed. Using a global dataset comprising 72 sites on six continents, we investigated which of 16 soil factors that shape nutrient availability associate most strongly with variation in grassland aboveground biomass. Climate and N deposition were also considered. Based on theory-driven structural equation modelling, we found that soil micronutrients (particularly Zn and Fe) were important predictors of biomass and, together with soil physicochemical properties and C:N, they explained more unique variation (32%) than climate and N deposition (24%). However, the association between micronutrients and biomass was absent in grasslands limited by NP. These results highlight soil properties as key predictors of global grassland biomass production and point to serial co-limitation by NP and micronutrients.

Universal scaling of robustness of ecosystem services to species loss.

Ensuring reliable supply of services from nature is key to the sustainable development and well-being of human societies. Varied and frequently complex relationships between biodiversity and ecosystem services have, however, frustrated our capacity to quantify and predict the vulnerability of those services to species extinctions. Here, we use a qualitative Boolean modelling framework to identify universal drivers of the robustness of ecosystem service supply to species loss. These drivers comprise simple features of the networks that link species to the functions they perform that, in turn, underpin a service. Together, they define what we call network fragility. Using data from >250 real ecological networks representing services such as pollination and seed-dispersal, we demonstrate that network fragility predicts remarkably well the robustness of empirical ecosystem services. We then show how to quantify contributions of individual species to ecosystem service robustness, enabling quantification of how vulnerability scales from species to services. Our findings provide general insights into the way species, functional traits, and the links between them together determine the vulnerability of ecosystem service supply to biodiversity loss.

General statistical scaling laws for stability in ecological systems.

Ecological stability refers to a family of concepts used to describe how systems of interacting species vary through time and respond to disturbances. Because observed ecological stability depends on sampling scales and environmental context, it is notoriously difficult to compare measurements across sites and systems. Here, we apply stochastic dynamical systems theory to derive general statistical scaling relationships across time, space, and ecological level of organisation for three fundamental stability aspects: resilience, resistance, and invariance. These relationships can be calibrated using random or representative samples measured at individual scales, and projected to predict average stability at other scales across a wide range of contexts. Moreover deviations between observed vs. extrapolated scaling relationships can reveal information about unobserved heterogeneity across time, space, or species. We anticipate that these methods will be useful for cross-study synthesis of stability data, extrapolating measurements to unobserved scales, and identifying underlying causes and consequences of heterogeneity.

Fertilized graminoids intensify negative drought effects on grassland productivity.

Droughts can strongly affect grassland productivity and biodiversity, but responses differ widely. Nutrient availability may be a critical factor explaining this variation, but is often ignored in analyses of drought responses. Here, we used a standardized nutrient addition experiment covering 10 European grasslands to test if full-factorial nitrogen, phosphorus, and potassium addition affected plant community responses to inter-annual variation in drought stress and to the extreme summer drought of 2018 in Europe. We found that nutrient addition amplified detrimental drought effects on community aboveground biomass production. Drought effects also differed between functional groups, with a negative effect on graminoid but not forb biomass production. Our results imply that eutrophication in grasslands, which promotes dominance of drought-sensitive graminoids over forbs, amplifies detrimental drought effects. In terms of climate change adaptation, agricultural management would benefit from taking into account differential drought impacts on fertilized versus unfertilized grasslands, which differ in ecosystem services they provide to society.

Opposing community assembly patterns for dominant and nondominant plant species in herbaceous ecosystems globally.

Biotic and abiotic factors interact with dominant plants-the locally most frequent or with the largest coverage-and nondominant plants differently, partially because dominant plants modify the environment where nondominant plants grow. For instance, if dominant plants compete strongly, they will deplete most resources, forcing nondominant plants into a narrower niche space. Conversely, if dominant plants are constrained by the environment, they might not exhaust available resources but instead may ameliorate environmental stressors that usually limit nondominants. Hence, the nature of interactions among nondominant species could be modified by dominant species. Furthermore, these differences could translate into a disparity in the phylogenetic relatedness among dominants compared to the relatedness among nondominants. By estimating phylogenetic dispersion in 78 grasslands across five continents, we found that dominant species were clustered (e.g., co-dominant grasses), suggesting dominant species are likely organized by environmental filtering, and that nondominant species were either randomly assembled or overdispersed. Traits showed similar trends for those sites (<50%) with sufficient trait data. Furthermore, several lineages scattered in the phylogeny had more nondominant species than expected at random, suggesting that traits common in nondominants are phylogenetically conserved and have evolved multiple times. We also explored environmental drivers of the dominant/nondominant disparity. We found different assembly patterns for dominants and nondominants, consistent with asymmetries in assembly mechanisms. Among the different postulated mechanisms, our results suggest two complementary hypotheses seldom explored: (1) Nondominant species include lineages adapted to thrive in the environment generated by dominant species. (2) Even when dominant species reduce resources to nondominant ones, dominant species could have a stronger positive effect on some nondominants by ameliorating environmental stressors affecting them, than by depleting resources and increasing the environmental stress to those nondominants. These results show that the dominant/nondominant asymmetry has ecological and evolutionary consequences fundamental to understand plant communities.

Individual species provide multifaceted contributions to the stability of ecosystems.

Exploration of the relationship between species diversity and ecological stability has occupied a prominent place in ecological research for decades. Yet, a key component of this puzzle-the contributions of individual species to the overall stability of ecosystems-remains largely unknown. Here, we show that individual species simultaneously stabilize and destabilize ecosystems along different dimensions of stability, and also that their contributions to functional (biomass) and compositional stability are largely independent. By simulating experimentally the extinction of three consumer species (the limpet Patella, the periwinkle Littorina and the topshell Gibbula) from a coastal rocky shore, we found that the capacity to predict the combined contribution of species to stability from the sum of their individual contributions varied among stability dimensions. This implies that the nature of the diversity-stability relationship depends upon the dimension of stability under consideration, and may be additive, synergistic or antagonistic. We conclude that, although the profoundly multifaceted and context-dependent consequences of species loss pose a significant challenge, the predictability of cumulative species contributions to some dimensions of stability provide a way forward for ecologists trying to conserve ecosystems and manage their stability under global change.

Warming can alter host behavior in a similar manner to infection with behavior-manipulating parasites.

Parasites are ecologically ubiquitous and, by modifying the physiology and behavior of their host organisms, act as key regulators of the dynamics and stability of ecosystems. It is, however, as yet unclear how parasitic relationships will act to moderate or accelerate the ecological impacts of global climate change. Here, we explore experimentally how the effects of parasites on both the physiology and behavior of their hosts can be moderated by warming, utilising a well-established aquatic host-parasite model system-the ecologically important amphipod Gammarus duebeni and its acanthocephalan parasite Polymorphus minutus. We show that, while only warming affected measured components of host physiology, parasite infection and warming both supressed predator-avoidance behavior of the host independently, yet in a similar manner. Six degrees of warming altered geotactic behaviors to the same extent as infection with behavior-manipulating parasites. These results indicate a novel mechanism by which parasites impact their ecosystems that could be critical to predicting the ecological impacts of warming. Our findings highlight the need for holistic knowledge of interaction networks, incorporating multiple interaction types and behaviors, to predict the effects of both warming and parasitism on the dynamics and stability of ecosystems.