Biodiversity research typically focuses on species richness and has often neglected interactions, either by assuming that such interactions are homogeneously distributed or by addressing only the interactions between a pair of species or a few species at a time. In contrast, a network approach provides a powerful representation of the ecological interactions among species and highlights their global interdependence. Understanding how the responses of pairwise interactions scale to entire assemblages remains one of the great challenges that must be met as society faces global ecosystem change.

In 2002, world leaders committed, through the Convention on Biological Diversity, to achieve a significant reduction in the rate of biodiversity loss by 2010. We compiled 31 indicators to report on progress toward this target. Most indicators of the state of biodiversity (covering species' population trends, extinction risk, habitat extent and condition, and community composition) showed declines, with no significant recent reductions in rate, whereas indicators of pressures on biodiversity (including resource consumption, invasive alien species, nitrogen pollution, overexploitation, and climate change impacts) showed increases. Despite some local successes and increasing responses (including extent and biodiversity coverage of protected areas, sustainable forest management, policy responses to invasive alien species, and biodiversity-related aid), the rate of biodiversity loss does not appear to be slowing.

Top predators often have powerful direct effects on prey populations, but whether these direct effects propagate to the base of terrestrial food webs is debated. There are few examples of trophic cascades strong enough to alter the abundance and composition of entire plant communities. We show that the introduction of arctic foxes (Alopex lagopus) to the Aleutian archipelago induced strong shifts in plant productivity and community structure via a previously unknown pathway. By preying on seabirds, foxes reduced nutrient transport from ocean to land, affecting soil fertility and transforming grasslands to dwarf shrub/forb-dominated ecosystems.

*The absence of co-evolved mutualists of plants invading a novel habitat is the logical corollary of the more widely recognized 'enemy escape'. To avoid or overcome the loss of mutualists, plants may co-invade with nonnative mutualists, form novel associations with native mutualists or form associations with native cosmopolitan mutualists, which are native but not novel to the invading plant. *We tested these hypotheses by contrasting the ectomycorrhizal fungal communities associated with invasive Pinus contorta in New Zealand with co-occurring endemic Nothofagus solandri var. cliffortioides. *Fungal communities on Pinus were species poor (14 ectomycorrhizal species) and dominated by nonnative (93%) and cosmopolitan fungi (7%). Nothofagus had a species-rich (98 species) fungal community dominated by native Cortinarius and two cosmopolitan fungi. *These results support co-invasion by mutualists rather than novel associations as an important mechanism by which plants avoid or overcome the loss of mutualists, consistent with invasional meltdown.

The introduction of invasive species, which often differ functionally from the components of the recipient community, generates ecological impacts that propagate along the food web. This review aims to determine how consistent the impacts of aquatic invasions are across taxa and habitats. To that end, we present a global meta-analysis from 151 publications (733 cases), covering a wide range of invaders (primary producers, filter collectors, omnivores and predators), resident aquatic community components (macrophytes, phytoplankton, zooplankton, benthic invertebrates and fish) and habitats (rivers, lakes and estuaries). Our synthesis suggests a strong negative influence of invasive species on the abundance of aquatic communities, particularly macrophytes, zooplankton and fish. In contrast, there was no general evidence for a decrease in species diversity in invaded habitats, suggesting a time lag between rapid abundance changes and local extinctions. Invaded habitats showed increased water turbidity, nitrogen and organic matter concentration, which are related to the capacity of invaders to transform habitats and increase eutrophication. The expansion of invasive macrophytes caused the largest decrease in fish abundance, the filtering activity of filter collectors depleted planktonic communities, omnivores (including both facultative and obligate herbivores) were responsible for the greatest decline in macrophyte abundance, and benthic invertebrates were most negatively affected by the introduction of new predators. These impacts were relatively consistent across habitats and experimental approaches. Based on our results, we propose a framework of positive and negative links between invasive species at four trophic positions and the five different components of recipient communities. This framework incorporates both direct biotic interactions (predation, competition, grazing) and indirect changes to the water physicochemical conditions mediated by invaders (habitat alteration). Considering the strong trophic links that characterize aquatic ecosystems, this framework is relevant to anticipate the far-reaching consequences of biological invasions on the structure and functionality of aquatic ecosystems. © 2015 John Wiley & Sons Ltd.

Tropical forests continue to be felled and fragmented around the world. A key question is how rapidly species disappear from forest fragments and how quickly humans must restore forest connectivity to minimize extinctions. We surveyed small mammals on forest islands in Chiew Larn Reservoir in Thailand 5 to 7 and 25 to 26 years after isolation and observed the near-total loss of native small mammals within 5 years from <10-hectare (ha) fragments and within 25 years from 10- to 56-ha fragments. Based on our results, we developed an island biogeographic model and estimated mean extinction half-life (50% of resident species disappearing) to be 13.9 years. These catastrophic extinctions were probably partly driven by an invasive rat species; such biotic invasions are becoming increasingly common in human-modified landscapes. Our results are thus particularly relevant to other fragmented forest landscapes and suggest that small fragments are potentially even more vulnerable to biodiversity loss than previously thought.

Through competition for pollinators, invasive plants may suppress native flora. Community-level studies provide an integrative assessment of invasion impacts and insights into factors that influence the vulnerability of different native species. We investigated effects of the nonnative herb Lythrum salicaria on pollination of native species in 14 fens of the eastern United States. We compared visitors per flower for 122 native plant species in invaded and uninvaded fens and incorporated a landscape-scale experiment, removing L. salicaria flowers from three of the invaded fens. Total flower densities were more than three times higher in invaded than uninvaded or removal sites when L. salicaria was blooming. Despite an increase in number of visitors with number of flowers per area, visitors per native flower declined with increasing numbers of flowers. Therefore, L. salicaria invasion depressed visitation to native flowers. In removal sites, visitation to native flowers was similar to uninvaded sites, confirming the observational results and also suggesting that invasion had not generated a persistent build-up of visitor populations. To study species-level impacts, we examined effects of invasion on visitors per flower for the 36 plant species flowering in both invaded and uninvaded fens. On average, the effect of invasion represented about a 20% reduction in visits per flower. We measured the influence of plant traits on vulnerability to L. salicaria invasion using meta-analysis. Bilaterally symmetrical flowers experienced stronger impacts on visitation, and similarity in flower color to L. salicaria weakly intensified competition with the invader for visitors. Finally, we assessed the reproductive consequences of competition with the invader in a dominant flowering shrub, Dasiphora fruticosa. Despite the negative effect of invasion on pollinator visitation in this species, pollen limitation of seed production was not stronger in invaded than in uninvaded sites, suggesting little impact of competition for pollinators on its population demography. Negative effects on pollination of native plants by this copiously flowering invader appeared to be mediated by increases in total flower density that were not matched by increases in pollinator density. The strength of impact was modulated across native species by their floral traits and reproductive ecology.© 2016 The Authors. Ecology, published by Wiley Periodicals, Inc., on behalf of the Ecological Society of America.

A fundamental goal of the study of ecology is to determine the drivers of habitat-forming vegetation, with much emphasis given to the relative importance to vegetation of "bottom-up" forces such as the role of nutrients and "top-down" forces such as the influence of herbivores and their predators. For coastal vegetation (e.g., kelp, seagrass, marsh, and mangroves) it has been well demonstrated that alterations to bottom-up forcing can cause major disturbances leading to loss of dominant vegetation. One such process is anthropogenic nutrient loading, which can lead to major changes in the abundance and species composition of primary producers, ultimately affecting important ecosystem services. In contrast, much less is known about the relative importance of apex predators on coastal vegetated ecosystems because most top predator populations have been depleted or lost completely. Here we provide evidence that an unusual four-level trophic cascade applies in one such system, whereby a top predator mitigates the bottom-up influences of nutrient loading. In a study of seagrass beds in an estuarine ecosystem exposed to extreme nutrient loading, we use a combination of a 50-y time series analysis, spatial comparisons, and mesocosm and field experiments to demonstrate that sea otters (Enhydra lutris) promote the growth and expansion of eelgrass (Zostera marina) through a trophic cascade, counteracting the negative effects of agriculturally induced nutrient loading. Our results add to a small but growing body of literature illustrating that significant interactions between bottom-up and top-down forces occur, in this case with consequences for the conservation of valued ecosystem services provided by seagrass.

生物入侵已对入侵区的生态环境、社会经济和人类健康造成了严重的威胁, 成为了21世纪五大全球性环境问题之一。本文回顾了2000年以来, 中国生物入侵研究领域尤其是入侵种的多样性与格局、入侵机制及生态学效应、管理与控制等方面所取得的重要进展, 讨论了需进一步加强研究的领域, 以期为进一步拓展该领域研究的广度和深度、为我国的生物入侵预警预防和科学治理提供参考。据初步研究, 中国的入侵种数量已达529种, 其中陆生植物、陆生无脊椎动物和微生物为主要入侵类群; 原产地以北美洲和南美洲为主; 经济发达和气候温暖湿润的东部和南部省份入侵态势明显较西部和北部省份严重; 随着中国经济的进一步发展, 生物入侵问题将可能更加严峻。外来种的成功入侵是其内禀优势、资源机遇和人为干扰共同作用的结果; 其中, 表型可塑性、适应性进化、天敌释放、种间互利或偏利共生和新化感作用等因素对入侵起到了关键作用。生物入侵已对中国土著生态系统的生物多样性和生态系统服务功能造成了严重影响, 打破了生态系统的固有平衡, 危害或威胁到中国的农林牧渔业生产、交通航运、环境、人类健康和公共设施安全。针对生物入侵的管理与控制, 中国加强了包括检测监测、风险分析、生物防治、扩散阻断、根治灭除和生态恢复等技术体系的研究和实施, 并初步控制了一些重要入侵种的扩张。中国生物入侵需要在全境性科学考察、生物入侵的遗传学、基因组学、生态系统影响、全球变化和管理与控制技术创新等领域进一步加强跨领域的交叉合作和系统研究。

Food web theory suggests that the placement of a weak interaction is critical such that under some conditions even one well-placed weak interaction can stabilise multiple strong interactions. This theory suggests that complex stable webs may be built from pivotal weak interactions such that the removal of even one to a few keystone interactions can have significant cascading impacts on whole system diversity and structure. However, the connection between weak interactions, derived from the theory of modular food web components, and keystone species, derived from empirical results, is not yet well understood. Here, we develop numerical techniques to detect potential oscillators hidden in complex food webs, and show that, both in random and real food webs, keystone consumer-resource interactions often operate to stabilise them. Alarmingly, this result suggests that nature frequently may be dangerously close to precipitous change with even the loss of one or a few weakly interacting species.© 2018 John Wiley & Sons Ltd/CNRS.

Ecology Letters (2010) 13: 1459-1474 ABSTRACT: There is growing concern that rapid environmental degradation threatens mutualistic interactions. Because mutualisms can bind species to a common fate, mutualism breakdown has the potential to expand and accelerate effects of global change on biodiversity loss and ecosystem disruption. The current focus on the ecological dynamics of mutualism under global change has skirted fundamental evolutionary issues. Here, we develop an evolutionary perspective on mutualism breakdown to complement the ecological perspective, by focusing on three processes: (1) shifts from mutualism to antagonism, (2) switches to novel partners and (3) mutualism abandonment. We then identify the evolutionary factors that may make particular classes of mutualisms especially susceptible or resistant to breakdown and discuss how communities harbouring mutualisms may be affected by these evolutionary responses. We propose a template for evolutionary research on mutualism resilience and identify conservation approaches that may help conserve targeted mutualisms in the face of environmental change.© 2010 Blackwell Publishing Ltd/CNRS.

Alien predators can have catastrophic effects on ecosystems and are thought to be much more harmful to biodiversity than their native counterparts. However, trophic cascade theory and the mesopredator release hypothesis predict that the removal of top predators will result in the reorganization of trophic webs and loss of biodiversity. Using field data collected throughout arid Australia, we provide evidence that removal of an alien top-predator, the dingo, has cascading effects through lower trophic levels. Dingo removal was linked to increased activity of herbivores and an invasive mesopredator, the red fox (Vulpes vulpes), and to the loss of grass cover and native species of small mammals. Using species distribution data, we predict that reintroducing or maintaining dingo populations would produce a net benefit for the conservation of threatened native mammals across greater than 2.42 x 10(6) km(2) of Australia. Our study provides evidence that an alien top predator can assume a keystone role and be beneficial for biodiversity conservation, and also that mammalian carnivores more generally can generate strong trophic cascades in terrestrial ecosystems.

Mutually beneficial interactions between flowering plants and animal pollinators represent a critical 'ecosystem service' under threat of anthropogenic extinction. We explored probable patterns of extinction in two large networks of plants and flower visitors by simulating the removal of pollinators and consequent loss of the plants that depend upon them for reproduction. For each network, we removed pollinators at random, systematically from least-linked (most specialized) to most-linked (most generalized), and systematically from most- to least-linked. Plant species diversity declined most rapidly with preferential removal of the most-linked pollinators, but declines were no worse than linear. This relative tolerance to extinction derives from redundancy in pollinators per plant and from nested topology of the networks. Tolerance in pollination networks contrasts with catastrophic declines reported from standard food webs. The discrepancy may be a result of the method used: previous studies removed species from multiple trophic levels based only on their linkage, whereas our preferential removal of pollinators reflects their greater risk of extinction relative to that of plants. In both pollination networks, the most-linked pollinators were bumble-bees and some solitary bees. These animals should receive special attention in efforts to conserve temperate pollination systems.

The assumption that climatic niche requirements of invasive species are conserved between their native and invaded ranges is key to predicting the risk of invasion. However, this assumption has been challenged recently by evidence of niche shifts in some species. Here, we report the first large-scale test of niche conservatism for 50 terrestrial plant invaders between Eurasia, North America, and Australia. We show that when analog climates are compared between regions, fewer than 15% of species have more than 10% of their invaded distribution outside their native climatic niche. These findings reveal that substantial niche shifts are rare in terrestrial plant invaders, providing support for an appropriate use of ecological niche models for the prediction of both biological invasions and responses to climate change.

The manner in which terrestrial ecosystems are regulated is controversial. The "top-down" school holds that predators limit herbivores and thereby prevent them from overexploiting vegetation. "Bottom-up" proponents stress the role of plant chemical defenses in limiting plant depredation by herbivores. A set of predator-free islands created by a hydroelectric impoundment in Venezuela allows a test of these competing world views. Limited area restricts the fauna of small (0.25 to 0.9 hectare) islands to predators of invertebrates (birds, lizards, anurans, and spiders), seed predators (rodents), and herbivores (howler monkeys, iguanas, and leaf-cutter ants). Predators of vertebrates are absent, and densities of rodents, howler monkeys, iguanas, and leaf-cutter ants are 10 to 100 times greater than on the nearby mainland, suggesting that predators normally limit their populations. The densities of seedlings and saplings of canopy trees are severely reduced on herbivore-affected islands, providing evidence of a trophic cascade unleashed in the absence of top-down regulation.

Efforts to understand the ecological regulation of species diversity via bottom-up approaches have failed to yield a consensus theory. Theories based on the alternative of top-down regulation have fared better. Paine's discovery of keystone predation demonstrated that the regulation of diversity via top-down forcing could be simple, strong, and direct, yet ecologists have persistently failed to perceive generality in Paine's result. Removing top predators destabilizes many systems and drives transitions to radically distinct alternative states. These transitions typically involve community reorganization and loss of diversity, implying that top-down forcing is crucial to diversity maintenance. Contrary to the expectations of bottom-up theories, many terrestrial herbivores and mesopredators are capable of sustained order-of-magnitude population increases following release from predation, negating the assumption that populations of primary consumers are resource limited and at or near carrying capacity. Predation sensu lato (to include Janzen-Connell mortality agents) has been shown to promote diversity in a wide range of ecosystems, including rocky intertidal shelves, coral reefs, the nearshore ocean, streams, lakes, temperate and tropical forests, and arctic tundra. The compelling variety of these ecosystems suggests that top-down forcing plays a universal role in regulating diversity. This conclusion is further supported by studies showing that the reduction or absence of predation leads to diversity loss and, in the more dramatic cases, to catastrophic regime change. Here, I expand on the thesis that diversity is maintained by the interaction between predation and competition, such that strong top-down forcing reduces competition, allowing coexistence.

Mutualisms structure ecosystems and mediate their functioning. They also enhance invasions of many alien species. Invasions disrupt native mutualisms, often leading to population declines, reduced biodiversity, and altered ecosystem functioning. Focusing on three main types of mutualisms (pollination, seed dispersal, and plant-microbial symbioses) and drawing on examples from different ecosystems and from species-and community-level studies, we review the key mechanisms whereby such positive interactions mediate invasions and are in turn influenced by invasions. High interaction generalization is "the norm" in most systems, allowing alien species to infiltrate recipient communities. We identify traits that influence invasiveness (e.g., selfing capacity in plants, animal behavioral traits) or invasibility (e.g., partner choice in mycorrhizas/rhizobia) through mutualistic interactions. Mutualistic disruptions due to invasions are pervasive, and subsequent cascading effects are also widespread. Ecological networks provide a useful framework for predicting tipping points for community collapse in response to invasions and other synergistic drivers of global change.

The main drivers of global environmental change (CO2 enrichment, nitrogen deposition, climate, biotic invasions and land use) cause extinctions and alter species distributions, and recent evidence shows that they exert pervasive impacts on various antagonistic and mutualistic interactions among species. In this review, we synthesize data from 688 published studies to show that these drivers often alter competitive interactions among plants and animals, exert multitrophic effects on the decomposer food web, increase intensity of pathogen infection, weaken mutualisms involving plants, and enhance herbivory while having variable effects on predation. A recurrent finding is that there is substantial variability among studies in both the magnitude and direction of effects of any given GEC driver on any given type of biotic interaction. Further, we show that higher order effects among multiple drivers acting simultaneously create challenges in predicting future responses to global environmental change, and that extrapolating these complex impacts across entire networks of species interactions yields unanticipated effects on ecosystems. Finally, we conclude that in order to reliably predict the effects of GEC on community and ecosystem processes, the greatest single challenge will be to determine how biotic and abiotic context alters the direction and magnitude of GEC effects on biotic interactions.

The formal protection of the Hawaiian green turtle Chelonia mydas in the 1970s has led to significant increases in the number of individuals recorded throughout the archipelago. Reduced growth rates and poor body condition of individuals at a number of foraging sites, including Kaloko-Honokohau National Historical Park (Kaloko), suggest that some aggregations have reached carrying capacity. To better understand the ecological structure and processes of the reef system at the park, we developed an ecosystem model that synthesized available data on Kaloko for the year 2005 and included 26 groups, spanning the entire trophic web. Model results showed that the combined grazing pressure of the different herbivore functional groups (i.e. reef fish, sea urchins, and green turtles) in Kaloko matched total algal production. Sea urchins exerted the strongest control over algal resources, partly because of their large biomass in park waters. Results confirmed that the Kaloko green turtle aggregation has reached carrying capacity. Green turtles help maintain low algal cover, and thus resilience of reefs in the face of disturbance, and should be explicitly included in studies of ecosystem dynamics on reefs. Our work also provides a 'current-condition' baseline for Kaloko, and a valuable tool for the assessment of the future marine ecosystem impacts of projected urban expansion plans around the park.

Ecosystems worldwide are losing some species and gaining others, resulting in an interchange of species that is having profound impacts on how these ecosystems function. However, research on the effects of species gains and losses has developed largely independently of one another. Recent conceptual advances regarding effects of species gain have arisen from studies that have unraveled the mechanistic basis of how invading species with novel traits alter biotic interactions and ecosystem processes. In contrast, studies on traits associated with species loss are fewer, and much remains unknown about how traits that predispose species to extinction affect ecological processes. Species gains and losses are both consequences and drivers of global change; thus, explicit integration of research on how both processes simultaneously affect ecosystem functioning is key to determining the response of the Earth system to current and future human activities.

Climate change research has demonstrated that changing temperatures will have an effect on community-level dynamics by altering species survival rates, shifting species distributions, and ultimately, creating mismatches in community interactions. However, most of this work has focused on increasing temperature, and still little is known about how the variation in temperature extremes will affect community dynamics. We used the model aquatic community held within the leaves of the carnivorous plant, Sarracenia purpurea, to test how food web dynamics will be affected by high temperature variation. We tested the community response of the first (bacterial density), second (protist diversity and composition), and third trophic level (predator mortality), and measured community respiration. We collected early and late successional stage inquiline communities from S. purpurea from two North American and two European sites with similar average July temperature. We then created a common garden experiment in which replicates of these communities underwent either high or normal daily temperature variation, with the average temperature equal among treatments. We found an impact of temperature variation on the first two, but not on the third trophic level. For bacteria in the high-variation treatment, density experienced an initial boost in growth but then decreased quickly through time. For protists in the high-variation treatment, alpha-diversity decreased faster than in the normal-variation treatment, beta-diversity increased only in the European sites, and protist community composition tended to diverge more in the late successional stage. The mortality of the predatory mosquito larvae was unaffected by temperature variation. Community respiration was lower in the high-variation treatment, indicating a lower ecosystem functioning. Our results highlight clear impacts of temperature variation. A more mechanistic understanding of the effects that temperature, and especially temperature variation, will have on community dynamics is still greatly needed.© 2016 John Wiley & Sons Ltd.

Phenology is a harbinger of climate change, with many species advancing flowering in response to rising temperatures. However, there is tremendous variation among species in phenological response to warming, and any phenological differences between native and non-native species may influence invasion outcomes under global warming. We simulated global warming in the field and found that non-native species flowered earlier and were more phenologically plastic to temperature than natives, which did not accelerate flowering in response to warming. Non-native species' flowering also became more synchronous with other community members under warming. Earlier flowering was associated with greater geographic spread of non-native species, implicating phenology as a potential trait associated with the successful establishment of non-native species across large geographic regions. Such phenological differences in both timing and plasticity between native and non-natives are hypothesised to promote invasion success and population persistence, potentially benefiting non-native over native species under climate change.© 2019 John Wiley & Sons Ltd/CNRS.

Species extinctions are accelerating globally, yet the mechanisms that maintain local biodiversity remain poorly understood. The extinction of species that feed on or are fed on by many others (i.e. 'hubs') has traditionally been thought to cause the greatest threat of further biodiversity loss. Very little attention has been paid to the strength of those feeding links (i.e. link weight) and the prevalence of indirect interactions. Here, we used a dynamical model based on empirical energy budget data to assess changes in ecosystem stability after simulating the loss of species according to various extinction scenarios. Link weight and/or indirect effects had stronger effects on food-web stability than the simple removal of 'hubs', demonstrating that both quantitative fluxes and species dissipating their effects across many links should be of great concern in biodiversity conservation, and the potential for 'hubs' to act as keystone species may have been exaggerated to date.© 2016 The Authors Ecology Letters published by CNRS and John Wiley & Sons Ltd.