Positive associations fuel soil biodiversity and ecological networks worldwide.

Microbial interactions are key to maintaining soil biodiversity. However, whether negative or positive associations govern the soil microbial system at a global scale remains virtually unknown, limiting our understanding of how microbes interact to support soil biodiversity and functions. Here, we explored ecological networks among multitrophic soil organisms involving bacteria, protists, fungi, and invertebrates in a global soil survey across 20 regions of the planet and found that positive associations among both pairs and triads of soil taxa governed global soil microbial networks. We further revealed that soil networks with greater levels of positive associations supported larger soil biodiversity and resulted in lower network fragility to withstand potential perturbations of species losses. Our study provides unique evidence of the widespread positive associations between soil organisms and their crucial role in maintaining the multitrophic structure of soil biodiversity worldwide.

Rapid monitoring of ecological persistence.

Effective conservation of ecological communities requires accurate and up-to-date information about whether species are persisting or declining to extinction. The persistence of an ecological community is supported by its underlying network of species interactions. While the persistence of the network supporting the whole community is the most relevant scale for conservation, in practice, only small subsets of these networks can be monitored. There is therefore an urgent need to establish links between the small snapshots of data conservationists can collect, and the "big picture" conclusions about ecosystem health demanded by policymakers, scientists, and societies. Here, we show that the persistence of small subnetworks (motifs) in isolation-that is, their persistence when considered separately from the larger network of which they are a part-is a reliable probabilistic indicator of the persistence of the network as a whole. Our methods show that it is easier to detect if an ecological community is not persistent than if it is persistent, allowing for rapid detection of extinction risk in endangered systems. Our results also justify the common practice of predicting ecological persistence from incomplete surveys by simulating the population dynamics of sampled subnetworks. Empirically, we show that our theoretical predictions are supported by data on invaded networks in restored and unrestored areas, even in the presence of environmental variability. Our work suggests that coordinated action to aggregate information from incomplete sampling can provide a means to rapidly assess the persistence of entire ecological networks and the expected success of restoration strategies.

A quantitative framework for identifying the role of individual species in Nature's Contributions to People.

It is widely acknowledged that biodiversity change is affecting human well-being by altering the supply of Nature's Contributions to People (NCP). Nevertheless, the role of individual species in this relationship remains obscure. In this article, we present a framework that combines the cascade model from ecosystem services research with network theory from community ecology. This allows us to quantitatively link NCP demanded by people to the networks of interacting species that underpin them. We show that this "network cascade" framework can reveal the number, identity and importance of the individual species that drive NCP and of the environmental conditions that support them. This information is highly valuable in demonstrating the importance of biodiversity in supporting human well-being and can help inform the management of biodiversity in social-ecological systems.

The Role of Indirect Effects in Coevolution along the Mutualism-Antagonism Continuum.

AbstractThe web of interactions in a community drives the coevolution of species. Yet it is unclear how the outcome of species interactions influences the coevolutionary dynamics of communities. This is a pressing matter, as changes to the outcome of interactions may become more common with human-induced global change. Here, we combine network and evolutionary theory to explore coevolutionary outcomes in communities harboring mutualistic and antagonistic interactions. We show that as the ratio of mutualistic to antagonistic interactions decreases, selection imposed by direct partners outweighs that imposed by indirect partners. This weakening of indirect effects results in communities composed of species with dissimilar traits and fast rates of adaptation. These changes are more pronounced when specialist consumers are the first species to engage in antagonistic interactions. Hence, a shift in the outcome of species interactions may reverberate across communities and alter the direction and speed of coevolution.

Dietary metabarcoding reveals the simplification of bird-pest interaction networks across a gradient of agricultural cover.

Agriculture is vital for supporting human populations, but its intensification often leads to landscape homogenization and a decline in non-provisioning ecosystem services. Ecological intensification and multifunctional landscapes are suggested as nature-based alternatives to intensive agriculture, using ecological processes like natural pest regulation to maximize food production. Birds are recognized for their role in increasing crop yields by consuming invertebrate pests in several agroecosystems. However, the understanding of how bird species, their traits and agricultural land cover influence the structure of bird-pest interactions remains limited. We sampled bird-pest interactions monthly for 1 year, at four sites within a multifunctional landscape, following a gradient of increasing agricultural land cover. We analysed 2583 droppings of 55 bird species with DNA metabarcoding and detected 225 pest species in 1139 samples of 42 bird species. As expected, bird-pest interactions were highly variable across bird species. Dietary pest richness was lower in the fully agricultural site, while predation frequency remained consistent across the agricultural land cover gradient. Network analysis revealed a reduction in the complexity of bird-pest interactions as agricultural coverage increased. Bird species abundance affected the bird's contribution to the network structure more than any of the bird traits analysed (weight, phenology, invertebrate frequency in diet and foraging strata), with more common birds being more important to network structure. Overall, our results show that increasing agricultural land cover increases the homogenization of bird-pest interactions. This shows the importance of maintaining natural patches within agricultural landscapes for biodiversity conservation and enhanced biocontrol.

A agricultura é essencial para suportar a população humana, mas a sua intensificação geralmente leva à homogeneização da paisagem e à redução dos serviços do ecossistema que não sejam de provisão. A intensificação ecológica e paisagens multifuncionais são sugeridas como alternativas naturais à agricultura intensiva, utilizando processos ecológicos como a regulação natural de pragas para maximizar a produção de alimentos. As aves são conhecidas pelo seu papel no aumento da produtividade das culturas por consumirem pragas em diversos agroecossistemas. Contudo, o conhecimento de como as espécies de aves, as suas características e a cobertura agrícola influenciam as interações entre aves e pragas são limitados. Nós amostrámos estas interações mensalmente durante um ano, em quatro locais, numa paisagem multifuncional, ao longo um gradiente de aumento da cobertura agrícola. Analisamos 2583 dejetos de 55 espécies de aves com DNA metabarcoding e detetamos 225 espécies praga em 1139 amostras de 42 espécies de aves. Como esperado, as interações entre aves e pragas foram muito distintas entre as várias espécies de aves. A riqueza de pragas na dieta foi menor no local completamente dominado por área agrícola, enquanto a frequência de predação de pragas foi constante ao longo do gradiente de cobertura agrícola. A análise de redes demonstrou uma redução na complexidade das interações entre aves e pragas à medida que a cobertura agrícola aumenta. A abundância das espécies de aves influenciou mais a contribuição das aves para a estrutura da rede do que qualquer uma das características analisadas (peso, fenologia, frequência de invertebrados na dieta e estrato de alimentação), sendo as aves mais comuns as mais importantes na estrutura da rede. De forma geral, os nossos resultados indicam que o aumento da cobertura agrícola aumenta a homogeneização das interações entre aves e pragas. Isto demonstra a importância de preservar áreas naturais em paisagem agrícolas para a conservação de biodiversidade e melhor controlo biológico.

Quantifying co-extinctions and ecosystem service vulnerability in coastal ecosystems experiencing climate warming.

Climate change is negatively impacting ecosystems and their contributions to human well-being, known as ecosystem services. Previous research has mainly focused on the direct effects of climate change on species and ecosystem services, leaving a gap in understanding the indirect impacts resulting from changes in species interactions within complex ecosystems. This knowledge gap is significant because the loss of a species in a food web can lead to additional species losses or "co-extinctions," particularly when the species most impacted by climate change are also the species that play critical roles in food web persistence or provide ecosystem services. Here, we present a framework to investigate the relationships among species vulnerability to climate change, their roles within the food web, their contributions to ecosystem services, and the overall persistence of these systems and services in the face of climate-induced species losses. To do this, we assess the robustness of food webs and their associated ecosystem services to climate-driven species extinctions in eight empirical rocky intertidal food webs. Across food webs, we find that highly connected species are not the most vulnerable to climate change. However, we find species that directly provide ecosystem services are more vulnerable to climate change and more connected than species that do not directly provide services, which results in ecosystem service provision collapsing before food webs. Overall, we find that food webs are more robust to climate change than the ecosystem services they provide and show that combining species roles in food webs and services with their vulnerability to climate change offer predictions about the impacts of co-extinctions for future food web and ecosystem service persistence. However, these conclusions are limited by data availability and quality, underscoring the need for more comprehensive data collection on linking species roles in interaction networks and their vulnerabilities to climate change.

Disturbance-mediated changes to boreal mammal spatial networks in industrializing landscapes.

Compound effects of anthropogenic disturbances on wildlife emerge through a complex network of direct responses and species interactions. Land-use changes driven by energy and forestry industries are known to disrupt predator-prey dynamics in boreal ecosystems, yet how these disturbance effects propagate across mammal communities remains uncertain. Using structural equation modeling, we tested disturbance-mediated pathways governing the spatial structure of multipredator multiprey boreal mammal networks across a landscape-scale disturbance gradient within Canada's Athabasca oil sands region. Linear disturbances had pervasive direct effects, increasing site use for all focal species, except black bears and threatened caribou, in at least one landscape. Conversely, block (polygonal) disturbance effects were negative but less common. Indirect disturbance effects were widespread and mediated by caribou avoidance of wolves, tracking of primary prey by subordinate predators, and intraguild dependencies among predators and large prey. Context-dependent responses to linear disturbances were most common among prey and within the landscape with intermediate disturbance. Our research suggests that industrial disturbances directly affect a suite of boreal mammals by altering forage availability and movement, leading to indirect effects across a range of interacting predators and prey, including the keystone snowshoe hare. The complexity of network-level direct and indirect disturbance effects reinforces calls for increased investment in addressing habitat degradation as the root cause of threatened species declines and broader ecosystem change.

Network motifs involving both competition and facilitation predict biodiversity in alpine plant communities.

Biological diversity depends on multiple, cooccurring ecological interactions. However, most studies focus on one interaction type at a time, leaving community ecologists unsure of how positive and negative associations among species combine to influence biodiversity patterns. Using surveys of plant populations in alpine communities worldwide, we explore patterns of positive and negative associations among triads of species (modules) and their relationship to local biodiversity. Three modules, each incorporating both positive and negative associations, were overrepresented, thus acting as "network motifs." Furthermore, the overrepresentation of these network motifs is positively linked to species diversity globally. A theoretical model illustrates that these network motifs, based on competition between facilitated species or facilitation between inferior competitors, increase local persistence. Our findings suggest that the interplay of competition and facilitation is crucial for maintaining biodiversity.

Are human-induced changes good or bad to dynamic landscape connectivity?

Land managers must find a compromise between rapidly changing landscapes and biodiversity conservation through ecological networks. Estimating ecological networks is a key approach to enhance or maintain functional connectivity by identifying the nodes and links of a graph, which represent habitats and their corresponding functional corridors, respectively. To understand the current state of biodiversity, it is necessary to consider dynamic landscape connectivity while relying on relevant land cover maps. Although a current land cover map is relatively easy to produce using existing data, this is challenging for past landscapes. Here we investigated the impact of changes in landscape connectivity in an urban landscape at a fine scale on the habitat availability of two bird species: the tree pipit Anthus trivialis and the short-toed treecreeper Certhia brachydactyla. These species, exhibiting different niche ecologies, have shown contrasting population trends at a medium-term scale. The occurrences of C. brachydactyla were better correlated with resistance values that maximise the use of corridors, whereas the occurrences of A. trivialis better fitted with intermediate resistance values. The statistical approach indirectly highlighted relevant information about the ecology the capacity of both species to use urban habitats. Landscape connectivity increased for both species over the 24-year study period and may have implications for local abundances, which could explain, at the national scale, the increase in populations of C. brachydactyla, but not the decrease in populations of A. trivialis. Thus, more attention must be paid on rural habitats and their associated species that are more impacted by human activities, but efforts could also be achieved in urban areas especially for highly corridor-dependent species. Studying dynamic landscape connectivity at a fine scale is essential for estimating past and future land cover changes and their associated impacts on ecological networks, to better reconcile human and biodiversity concerns in land management.

Using simulation modeling to demonstrate the performance of graph theory metrics and connectivity algorithms.

Models and metrics to measure ecological connectivity are now well-developed and widely used in research and applications to mitigate the ecological impacts of climate change and anthropogenic habitat loss. Despite the prevalent application of connectivity models, however, relatively little is known about the performance of these methods in predicting functional connectivity patterns and organism movement. Our goal in this paper was to compare different connectivity models in their abilities to predict a wide range of simulated animal movement patterns. We used the Pathwalker software to evaluate the performance of several connectivity model predictions based on graph theory, resistant kernels, and factorial least-cost paths. In addition, we assessed the efficacy of synoptic and patch-based approaches to defining source points for analysis. In total, we produced 28 different simulations of animal movement. As we expected, we found that the choice of connectivity model used was the variable that most influenced prediction accuracy. Moreover, we found that the resistant kernels approach consistently provided the strongest correlations to the simulated underlying movement processes. The results also suggested that the agent-based simulation approach itself can often be the best analytical framework to map functional connectivity for ecological research and conservation applications, given its biological realism and flexibility to implement combinations of movement mechanism, dispersal threshold, directional bias, destination bias and spatial composition of source locations for analysis. In doing so, we provide novel insights to guide future functional connectivity analyses. In future research, we could use the same model for several different species groups and see how this reliability depends on the species analyzed. This could bring to light other elements that play an essential role in predicting connectivity.

Five fundamental ways in which complex food webs may spiral out of control.

Theory suggests that increasingly long, negative feedback loops of many interacting species may destabilize food webs as complexity increases. Less attention has, however, been paid to the specific ways in which these 'delayed negative feedbacks' may affect the response of complex ecosystems to global environmental change. Here, we describe five fundamental ways in which these feedbacks might pave the way for abrupt, large-scale transitions and species losses. By combining topological and bioenergetic models, we then proceed by showing that the likelihood of such transitions increases with the number of interacting species and/or when the combined effects of stabilizing network patterns approach the minimum required for stable coexistence. Our findings thus shift the question from the classical question of what makes complex, unaltered ecosystems stable to whether the effects of, known and unknown, stabilizing food-web patterns are sufficient to prevent abrupt, large-scale transitions under global environmental change.

Reciprocity and interaction effectiveness in generalised mutualisms among free-living species.

Mutualistic interactions among free-living species generally involve low-frequency interactions and highly asymmetric dependence among partners, yet our understanding of factors behind their emergence is still limited. Using individual-based interactions of a super-generalist fleshy-fruited plant with its frugivore assemblage, we estimated the Resource Provisioning Effectiveness (RPE) and Seed Dispersal Effectiveness (SDE) to assess the balance in the exchange of resources. Plants were highly dependent on a few frugivore species, while frugivores interacted with most individual plants, resulting in strong asymmetries of mutual dependence. Interaction effectiveness was mainly driven by interaction frequency. Despite highly asymmetric dependences, the strong reliance on quantity of fruit consumed determined high reciprocity in rewards between partners (i.e. higher energy provided by the plant, more seedlings recruited), which was not obscured by minor variations in the quality of animal or plant service. We anticipate reciprocity will emerge in low-intimacy mutualisms where the mutualistic outcome largely relies upon interaction frequency.

Decision-making under uncertainty for species introductions into ecological networks.

Ecological communities are increasingly subject to natural and human-induced additions of species, as species shift their ranges under climate change, are introduced for conservation and are unintentionally moved by humans. As such, decisions about how to manage ecosystems subject to species introductions and considering multiple management objectives need to be made. However, the impacts of gaining new species on ecological communities are difficult to predict due to uncertainty in introduced species characteristics, the novel interactions that will be produced by that species, and the recipient ecosystem structure. Drawing on ecological and conservation decision theory, we synthesise literature into a conceptual framework for species introduction decision-making based on ecological networks in high-uncertainty contexts. We demonstrate the application of this framework to a theoretical decision surrounding assisted migration considering both biodiversity and ecosystem service objectives. We show that this framework can be used to evaluate trade-offs between outcomes, predict worst-case scenarios, suggest when one should collect additional data, and allow for improving knowledge of the system over time.

Invasive species modulate the structure and stability of a multilayer mutualistic network.

Species interactions are critical for maintaining community structure and dynamics, but the effects of invasive species on multitrophic networks remain poorly understood. We leveraged an ongoing invasion scenario in Patagonia, Argentina, to explore how non-native ungulates affect multitrophic networks. Ungulates disrupt a hummingbird-mistletoe-marsupial keystone interaction, which alters community composition. We sampled pollination and seed dispersal interactions in intact and invaded sites. We constructed pollination and seed dispersal networks for each site, which we connected via shared plants. We calculated pollination-seed dispersal connectivity, identified clusters of highly connected species, and quantified species' roles in connecting species clusters. To link structural variation to stability, we quantified network tolerance to single random species removal (disturbance propagation) and sequential species removal (robustness) using a stochastic coextinction model. Ungulates reduced the connectivity between pollination and seed dispersal and produced fewer clusters with a skewed size distribution. Moreover, species shifted their structural role, fragmenting the network by reducing the 'bridges' among species clusters. These structural changes altered the dynamics of cascading effects, increasing disturbance propagation and reducing network robustness. Our results highlight invasive species' role in altering community structure and subsequent stability in multitrophic communities.

The turnover of plant-frugivore interactions along plant range expansion: consequences for natural colonization processes.

Plant-animal mutualisms such as seed dispersal are key interactions for sustaining plant range shifts. It remains elusive whether the organization of interactions with seed dispersers is reconfigured along the expansion landscape template and, if so, whether its effects accelerate or slow colonization. Here we analyse plant-frugivore interactions in a scenario of rapid population expansion of a Mediterranean juniper. We combined network analyses with field surveys, sampling interactions between individual plants and frugivores by DNA-barcoding and phototrapping over two seasons. We assess the role of intrinsic and extrinsic intraspecific variability in shaping interactions and we estimate the individual plant contributions to the seed rain. The whole interaction network was highly structured, with a distinct set of modules including individual plants and frugivore species arranged concordantly along the expansion gradient. The modular configuration was partially shaped by individual neighbourhood context (density and fecundity) and phenotypic traits (cone size). Interaction reconfiguration resulted in a higher and more uneven propagule contribution, with most effective dispersers having a prominent role at the colonization front stand, where a distinct subset of early arriving plants dominated the seed rain. Our study offers new insights into the key role of mutualistic interactions in colonization scenarios by promoting fast plant expansion processes.

The reliability of regional ecological knowledge to build local interaction networks: a test using seed-dispersal networks across land-bridge islands.

Building ecological networks is the fundamental basis of depicting how species in communities interact, but sampling complex interaction networks is extremely labour intensive. Recently, indirect ecological information has been applied to build interaction networks. Here we propose to extend the source of indirect ecological information, and applied regional ecological knowledge to build local interaction networks. Using a high-resolution dataset consisting of 22 locally observed networks with 17 572 seed-dispersal events, we test the reliability of indirectly derived local networks based on regional ecological knowledge (REK) across islands. We found that species richness strongly influenced 'local interaction rewiring' (i.e. the proportion of locally observed interactions among regionally interacting species), and all network properties were biased using REK-based networks. Notably, species richness and local interaction rewiring strongly affected estimations of REK-based network structures. However, locally observed and REK-based networks detected the same trends of how network structure correlates to island area and isolation. These results suggest that we should use REK-based networks cautiously for reflecting actual interaction patterns of local networks, but highlight that REK-based networks have great potential for comparative studies across environmental gradients. The use of indirect regional ecological information may thus advance our understanding of biogeographical patterns of species interactions.

Ecological network structure in response to community assembly processes over evolutionary time.

The dynamic structure of ecological communities results from interactions among taxa that change with shifts in species composition in space and time. However, our ability to study the interplay of ecological and evolutionary processes on community assembly remains relatively unexplored due to the difficulty of measuring community structure over long temporal scales. Here, we made use of a geological chronosequence across the Hawaiian Islands, representing 50 years to 4.15 million years of ecosystem development, to sample 11 communities of arthropods and their associated plant taxa using semiquantitative DNA metabarcoding. We then examined how ecological communities changed with community age by calculating quantitative network statistics for bipartite networks of arthropod-plant associations. The average number of interactions per species (linkage density), ratio of plant to arthropod species (vulnerability) and uniformity of energy flow (interaction evenness) increased significantly in concert with community age. The index of specialization H 2 ' has a curvilinear relationship with community age. Our analyses suggest that younger communities are characterized by fewer but stronger interactions, while biotic associations become more even and diverse as communities mature. These shifts in structure became especially prominent on East Maui (~0.5 million years old) and older volcanos, after enough time had elapsed for adaptation and specialization to act on populations in situ. Such natural progression of specialization during community assembly is probably impeded by the rapid infiltration of non-native species, with special risk to younger or more recently disturbed communities that are composed of fewer specialized relationships.

Interkingdom ecological networks between plants and fungi drive soil multifunctionality across arid inland river basin.

Despite the known collective contribution of above- (plants) and below-ground (soil fungi) biodiversity on multiple soil functions, how the associations among plant and fungal communities regulate soil multifunctionality (SMF) differentially remains unknown. Here, plant communities were investigated at 81 plots across a typical arid inland river basin, within which associated soil fungal communities and seven soil functions (nutrients storage and biological activity) were measured in surface (0-15 cm) and subsurface soil (15-30 cm). We evaluated the relative importance of species richness and biotic associations (reflected by network complexity) on SMF. Our results demonstrated that plant species richness and plant-fungus network complexity promoted SMF in surface and subsurface soil. SMF in two soil layers was mainly determined by plant-fungus network complexity, mean groundwater depth and soil variables, among which plant-fungus network complexity played a crucial role. Plant-fungus network complexity had stronger effects on SMF in surface soil than in subsurface soil. We present evidence that plant-fungus network complexity surpassed plant-fungal species richness in determining SMF in surface and subsurface soil. Moreover, plant-fungal species richness could not directly affect SMF. Greater plant-fungal species richness indirectly promoted SMF since they ensured greater plant-fungal associations. Collectively, we concluded that interkingdom networks between plants and fungi drive SMF even in different soil layers. Our findings enhanced our knowledge of the underlying mechanisms that above- and below-ground associations promote SMF in arid inland river basins. Future study should place more emphasis on the associations among plant and microbial communities in protecting soil functions under global changes.

Temperature-mediated microbial carbon utilization in China's lakes.

Microbes play an important role in aquatic carbon cycling but we have a limited understanding of their functional responses to changes in temperature across large geographic areas. Here, we explored how microbial communities utilized different carbon substrates and the underlying ecological mechanisms along a space-for-time substitution temperature gradient of future climate change. The gradient included 47 lakes from five major lake regions in China spanning a difference of nearly 15°C in mean annual temperatures (MAT). Our results indicated that lakes from warmer regions generally had lower values of variables related to carbon concentrations and greater carbon utilization than those from colder regions. The greater utilization of carbon substrates under higher temperatures could be attributed to changes in bacterial community composition, with a greater abundance of Cyanobacteria and Actinobacteriota and less Proteobacteria in warmer lake regions. We also found that the core species in microbial networks changed with increasing temperature, from Hydrogenophaga and Rhodobacteraceae, which inhibited the utilization of amino acids and carbohydrates, to the CL500-29-marine-group, which promoted the utilization of all almost carbon substrates. Overall, our findings suggest that temperature can mediate aquatic carbon utilization by changing the interactions between bacteria and individual carbon substrates, and the discovery of core species that affect carbon utilization provides insight into potential carbon sequestration within inland water bodies under future climate warming.

Climate change and commercial fishing practices codetermine survival of a long-lived seabird.

Understanding the environmental mechanisms that govern population change is a fundamental objective in ecology. Although the determination of how top-down and bottom-up drivers affect demography is important, it is often equally critical to understand the extent to which, environmental conditions that underpin these drivers fluctuate across time. For example, associations between climate and both food availability and predation risk may suggest the presence of trophic interactions that may influence inferences made from patterns in ecological data. Analytical tools have been developed to account for these correlations, while providing opportunities to ask novel questions regarding how populations change across space and time. Here, we combine two modeling disciplines-path analysis and mark-recapture-recovery models-to explore whether shifts in sea-surface temperatures (SSTs) influenced top-down (entanglement in fishing equipment) or bottom-up (forage fish production) population constraints over 60 years, and the extent to which these covarying processes shaped the survival of a long-lived seabird, the Royal tern. We found that hemispheric trends in SST were associated with variation in the amount of fish harvested along the Atlantic coast of North America and in the Caribbean, whereas reductions in forage fish production were mostly driven by shifts in the amount of fish harvested by commercial fisheries throughout the North Atlantic the year prior. Although the indirect (i.e., stock depletion) and direct (i.e., entanglement) impacts of commercial fishing on Royal tern mortality has declined over the last 60 years, increased SSTs during this time period has resulted in a comparable increase in mortality risk, which disproportionately impacted the survival of the youngest age-classes of Royal terns. Given climate projections for the North Atlantic, our results indicate that threats to Royal tern population persistence in the Mid-Atlantic will most likely be driven by failures to recruit juveniles into the breeding population.