Expansion of corals on temperate reefs: direct and indirect effects of marine heatwaves

Globally, many temperate marine communities have experienced significant temperature increases over recent decades in the form of gradual warming and heatwaves. As a result, these communities are shifting towards increasingly subtropical and tropical species compositions. Expanding coral populations have been reported from several temperate reef ecosystems along warming coastlines; these changes have been attributed to direct effects of gradual warming over decades. In contrast, increases in coral populations following shorter-term extreme warming events have rarely been documented. In this study, we compared coral populations on 17 temperate reefs in Western Australia before (2005/06) and after (2013) multiple marine heatwaves (2010–2012) affected the entire coastline. We hypothesised that coral communities would expand and change as a consequence of increasing local populations and recruitment of warm-affinity species. We found differences in coral community structure over time, driven primarily by a fourfold increase of one local species, Plesiastrea versipora, rather than recruitment of warm-affinity species. Coral populations became strongly dominated by small size classes, indicative of recent increased recruitment or recruit survival. These changes were likely facilitated by competitive release of corals from dominant temperate seaweeds, which perished during the heatwaves, rather than driven by direct temperature effects. Overall, as corals are inherently warm-water taxa not commonly associated with seaweed-dominated temperate reefs, these findings are consistent with a net tropicalisation. Our study draws attention to processes other than gradual warming that also influence the trajectory of temperate reefs in a changing ocean.

     

Temperature and trophic structure are driving microbial productivity along a biogeographical gradient

Temperature is known to influence ecosystem processes through its direct effect on biological rates such as respiration and nutrient cycling. These changes can then indirectly affect ecologically processes by altering trophic dynamics, the persistence of a species in a given environment, and, consequently, its distribution. However, it is not known if this direct effect of temperature on biological rates is singularly the most important factor for the functioning of ecosystems, or if trophic structure and the adaptation of a species to the local environment also play an essential role. Understanding the relative importance of these factors is crucial for predicting the impact that climate change will have on species and ecosystems. To achieve a more complete understanding of the impact of changing temperatures, it is necessary to integrate perspectives from biogeography, such as the influences of species distribution and local adaptation, with ecosystem and community ecology. By using the microbial community inhabiting the water‐filled leaves of Sarracenia purpurea, we tested the importance of temperature, trophic structure, and local adaptation on ecosystem functioning. We accomplished this by collecting communities along a natural temperature gradient and maintaining these communities in a common garden, factorial experiment. To test for the importance of local adaptation and temperature, the origin of each community was crossed with the temperature from each site. Additionally, to test the importance of top‐down trophic regulation for ecosystem functioning, the presence of the mosquito larvae top predator was manipulated. We found that temperature has a greater effect on ecosystem functioning than origin, and that top‐down trophic regulation increased with temperature. Our results emphasize the synergistic effects of temperature and biotic interactions when predicting the consequences of global warming on ecosystem functioning.     

陆地生态系统凋落物分解对全球气候变暖的响应

陆地生态系统凋落物分解是全球碳收支的一个重要组成部分, 主要受气候、凋落物质量和土壤生物群落的综合控制。科学家们普遍认为全球气候变化将对陆地生态系统凋落物分解产生复杂而深远的影响。该文结合凋落物分解试验的常用方法——缩微试验、原位模拟实验和自然环境梯度实验, 归纳现有研究结果, 意在揭示全球气候变化对陆地生态系统凋落物分解的直接影响(温度对凋落物分解速率的影响)和间接影响(温度对凋落物质量、土壤微生物群落及植被型的影响)的普遍规律。各种研究方法都表明: 在水分条件理想的情况下, 温度升高往往能加快凋落物的分解速率; 原位模拟实验中, 凋落物分解速率因物种、增温方法和地理方位而异; 全球气候变化能改变凋落物质量, 但可能不会在短期内影响凋落物的分解速率; 凋落物质量和可分解性的种间差异远大于增温所引发的表型响应差异, 那么, 气候变化所引发的植物群落结构和物种组成的变化将对陆地生态系统凋落物分解产生更强烈的影响; 土壤生物群落如何响应全球气候变化, 进而怎样影响凋落物分解过程, 这些都还存在着极大的不确定性。     

Interactive effects of warming,eutrophication and size structure: impacts on biodiversity and food‐web structure

Warming and eutrophication are two of the most important global change stressors for natural ecosystems, but their interaction is poorly understood. We used a dynamic model of complex, size‐structured food webs to assess interactive effects on diversity and network structure. We found antagonistic impacts: Warming increases diversity in eutrophic systems and decreases it in oligotrophic systems. These effects interact with the community size structure: Communities of similarly sized species such as parasitoid–host systems are stabilized by warming and destabilized by eutrophication, whereas the diversity of size‐structured predator–prey networks decreases strongly with warming, but decreases only weakly with eutrophication. Nonrandom extinction risks for generalists and specialists lead to higher connectance in networks without size structure and lower connectance in size‐structured communities. Overall, our results unravel interactive impacts of warming and eutrophication and suggest that size structure may serve as an important proxy for predicting the community sensitivity to these global change stressors.     

Five Years of Experimental Warming Increases the Biodiversity and Productivity of Phytoplankton

Phytoplankton are key components of aquatic ecosystems, fixing CO₂ from the atmosphere through photosynthesis and supporting secondary production, yet relatively little is known about how future global warming might alter their biodiversity and associated ecosystem functioning. Here, we explore how the structure, function, and biodiversity of a planktonic metacommunity was altered after five years of experimental warming. Our outdoor mesocosm experiment was open to natural dispersal from the regional species pool, allowing us to explore the effects of experimental warming in the context of metacommunity dynamics. Warming of 4°C led to a 67% increase in the species richness of the phytoplankton, more evenly-distributed abundance, and higher rates of gross primary productivity. Warming elevated productivity indirectly, by increasing the biodiversity and biomass of the local phytoplankton communities. Warming also systematically shifted the taxonomic and functional trait composition of the phytoplankton, favoring large, colonial, inedible phytoplankton taxa, suggesting stronger top-down control, mediated by zooplankton grazing played an important role. Overall, our findings suggest that temperature can modulate species coexistence, and through such mechanisms, global warming could, in some cases, increase the species richness and productivity of phytoplankton communities.     

Subsidies mediate interactions between communities across space

Most spatial ecology focuses on how species dispersal affects community dynamics and coexistence. Ecosystems, however, are also commonly connected by flows of resources. We experimentally tested how neighbouring communities indirectly influence each other in absence of dispersal, via resource exchanges. Using two‐patch microcosm meta‐ecosystems, we manipulated community composition and dynamics, by varying separately species key functional traits (autotroph versus heterotroph species and size of consumer species) and trophic structure of aquatic communities (species growing alone or in presence of competitors or predators). We then analysed the effects of species functional traits and trophic structure on communities connected through spatial subsidies in the absence of actual dispersal. Both functional traits and trophic structure strongly affected dynamics across neighbouring communities. Heterotroph communities connected to autotroph neighbours developed better than with heterotroph neighbours, such that coexistence of competitors was determined by the functional traits of the neighbouring community. Densities in autotroph communities were also strikingly higher when receiving subsidies from heterotroph communities compared to their own subsidies when grown in isolated ecosystems. In contrast, communities connected to predator‐dominated ecosystems collapsed, without any direct contact with the predators. Our results demonstrate that because community composition and structure modify the distribution of biomass within a community, they may also affect communities connected through subsidies through quantitative and qualitative changes of detritus flows. This stresses that ecosystem management should account for such interdependencies mediated by spatial subsidies, given that local community alterations cascade across space onto other ecosystems even if species dispersal is completely absent.     

Temperature‐size responses alter food chain persistence across environmental gradients

Body‐size reduction is a ubiquitous response to global warming alongside changes in species phenology and distributions. However, ecological consequences of temperature‐size (TS) responses for community persistence under environmental change remain largely unexplored. Here, we investigated the interactive effects of warming, enrichment, community size structure and TS responses on a three‐species food chain using a temperature‐dependent model with empirical parameterisation. We found that TS responses often increase community persistence, mainly by modifying consumer‐resource size ratios and thereby altering interaction strengths and energetic efficiencies. However, the sign and magnitude of these effects vary with warming and enrichment levels, TS responses of constituent species, and community size structure. We predict that the consequences of TS responses are stronger in aquatic than in terrestrial ecosystems, especially when species show different TS responses. We conclude that considering the links between phenotypic plasticity, environmental drivers and species interactions is crucial to better predict global change impacts on ecosystem diversity and stability.     

Warming will affect phytoplankton differently: evidence through a mechanistic approach

Although the consequences of global warming in aquatic ecosystems are only beginning to be revealed, a key to forecasting the impact on aquatic communities is an understanding of individual species' vulnerability to increased temperature. Despite their microscopic size, phytoplankton support about half of the global primary production, drive essential biogeochemical cycles and represent the basis of the aquatic food web. At present, it is known that phytoplankton are important targets and, consequently, harbingers of climate change in aquatic systems. Therefore, investigating the capacity of phytoplankton to adapt to the predicted warming has become a relevant issue. However, considering the polyphyletic complexity of the phytoplankton community, different responses to increased temperature are expected. We experimentally tested the effects of warming on 12 species of phytoplankton isolated from a variety of environments by using a mechanistic approach able to assess evolutionary adaptation (the so-called ratchet technique). We found different degrees of tolerance to temperature rises and an interspecific capacity for genetic adaptation. The thermal resistance level reached by each species is discussed in relation to their respective original habitats. Our study additionally provides evidence on the most resistant phytoplankton groups in a future warming scenario.     

Size‐based ecological interactions drive food web responses to climate warming

Predicting climate change impacts on animal communities requires knowledge of how physiological effects are mediated by ecological interactions. Food‐dependent growth and within‐species size variation depend on temperature and affect community dynamics through feedbacks between individual performance and population size structure. Still, we know little about how warming affects these feedbacks. Using a dynamic stage‐structured biomass model with food‐, size‐ and temperature‐dependent life history processes, we analyse how temperature affects coexistence, stability and size structure in a tri‐trophic food chain, and find that warming effects on community stability depend on ecological interactions. Predator biomass densities generally decline with warming – gradually or through collapses – depending on which consumer life stage predators feed on. Collapses occur when warming induces alternative stable states via Allee effects. This suggests that predator persistence in warmer climates may be lower than previously acknowledged and that effects of warming on food web stability largely depend on species interactions.     

冬季升温对高山生态系统碳氮循环过程的影响

全球温度升高是目前面临的重要环境问题,但存在明显的季节差异性,即冬季升温幅度显著高于夏季的季节非对称性趋势,这在高纬度和高海拔地区更加显著。冬季升温会直接影响积雪覆盖与冰冻层厚度,并引起冻融交替循环的增加,而冬季植物处于休眠状态,这会直接影响土壤中有效氮的吸收与损失,引起土壤有效氮可利用性的变化。然而,关于冬季增温对后续生长季节植物活动、土壤碳氮循环过程的影响等方面的研究仍存在诸多不确定。综述了冬季升温对积雪覆盖与冻融交替循环改变对高山生态系统物质循环的影响,以及冬季升温对土壤碳氮循环、微生物与酶活性的影响,并由此引起的植物物候期、群落结构、生产与养分循环与凋落物分解等生理、生态过程方面的研究进展。在未来的研究中,应针对不同生态系统特点选择合适的冬季增温方式,加强非极地苔原地区关于冬季升温的研究,注重关注冬季升温对植物-土壤微生物之间反馈作用的影响,重点关注冬季升温对生态系统的延滞效应。     

Size‐balanced community reorganization in response to nutrients and warming

It is widely accepted that global warming will adversely affect ecological communities. As ecosystems are simultaneously exposed to other anthropogenic influences, it is important to address the effects of climate change in the context of many stressors. Nutrient enrichment might offset some of the energy demands that warming can exert on organisms by stimulating growth at the base of the food web. It is important to know whether indirect effects of warming will be as ecologically significant as direct physiological effects. Declining body size is increasingly viewed as a universal response to warming, with the potential to alter trophic interactions. To address these issues, we used an outdoor array of marine mesocosms to examine the impacts of warming, nutrient enrichment and altered top‐predator body size on a community comprised of the predator (shore crab Carcinus maenas), various grazing detritivores (amphipods) and algal resources. Warming increased mortality rates of crabs, but had no effect on their moulting rates. Nutrient enrichment and warming had near diametrically opposed effects on the assemblage, confirming that the ecological effects of these two stressors can cancel each other out. This suggests that nutrient‐enriched systems might act as an energy refuge to populations of species under metabolic constraints due to warming. While there was a strong difference in assemblages between mesocosms containing crabs compared to mesocosms without crabs, decreasing crab size had no detectable effect on the amphipod or algal assemblages. This suggests that in allometrically balanced communities, the expected long‐term effect of warming (declining body size) is not of similar ecological consequence to the direct physiological effects of warming, at least not over the six week duration of the experiment described here. More research is needed to determine the long‐term effects of declining body size on the bioenergetic balance of natural communities.     

Interactive effects of global warming and ‘global worming’ on the initial establishment of native and exotic herbaceous plant species

The spread of exotic earthworms (‘worming’) and rising temperatures are expected to alter the biological, chemical and physical properties of many ecosystems, yet little is known about their potential interactive effects. We performed a laboratory microcosm experiment to investigate the effects of earthworms (anecic, endogeic, epigeic, or all three together) and 4°C warming on soil water content, litter turnover and seedling establishment of four native and four exotic herbaceous plant species. Warming and worming exerted independent as well as interactive effects on soil processes and plant dynamics. Warming reduced the water content of the upper soil layer, but only in the presence of earthworms. Litter removal increased in the presence of earthworms, the effect being most pronounced in the presence of anecic earthworms at ambient temperature. Exotic plant species were most influenced by earthworms (lower seedling number but higher biomass), whereas natives were most sensitive to warming (higher seedling number). This differential response resulted in significant interaction effects of earthworms and warming on abundance and richness of native relative to exotic plants as well as related shifts in plant species composition. Structural equation modeling allowed us to address possible mechanisms: direct effects of earthworms primarily affected exotic plants, whereas earthworms and warming indirectly and differentially affected native and exotic plants through changes in soil water content and surface litter. Invasive earthworms and warming are likely to interactively impact abiotic and biotic ecosystem properties. The invasion of epigeic and anecic species could select for plant species able to germinate on bare soil and tolerate drought, with the latter becoming more important in a warmer world. Thus earthworm invasion may result in simplified plant communities of increased susceptibility to the invasion of exotic plants.     

Community monopolization: local adaptation enhances priority effects in an evolving metacommunity

The diversity and composition of biological communities might often depend on colonization history because early colonists can exclude future colonists through a priority effect. These priority effects, which have been observed across a wide variety of ecosystems, often arise because early colonists have sufficient time to use available resources efficiently and subsequently withhold them from invaders. Here, we explore the extent to which rapid local adaptive evolution contributes to the pervasiveness of these priority effects. Using an individual-based simulation, we show that early colonization allows the descendants of colonists to adapt to novel conditions and reduce the establishment success of an initially ecologically equivalent competing species. Our model predicts that slight differences in colonization timing and adaptive capacity between species can substantially alter the dynamics and diversity of communities. We also show that priority effects and gene flow can generate a novel mechanism for the expansion and retraction of species distributions in a metacommunity. Our results suggest that local adaptation combined with stochastic colonization events can obscure direct relationships between species distributions and environmental gradients. Given the increasing recognition of rapid, microgeographic evolution in natural populations, we expect that evolutionary priority effects could affect the structure and dynamics of many natural metacommunities.     

Plant Species Composition and Productivity following Permafrost Thaw and Thermokarst in Alaskan Tundra

Climate warming is expected to have a large impact on plant species composition and productivity in northern latitude ecosystems. Warming can affect vegetation communities directly through temperature effects on plant growth and indirectly through alteration of soil nutrient availability. In addition, warming can cause permafrost to thaw and thermokarst (ground subsidence) to develop, which can alter the structure of the ecosystem by altering hydrological patterns within a site. These multiple direct and indirect effects of permafrost thawing are difficult to simulate in experimental approaches that often manipulate only one or two factors. Here, we used a natural gradient approach with three sites to represent stages in the process of permafrost thawing and thermokarst. We found that vascular plant biomass shifted from graminoid-dominated tundra in the least disturbed site to shrub-dominated tundra at the oldest, most subsided site, whereas the intermediate site was co-dominated by both plant functional groups. Vascular plant productivity patterns followed the changes in biomass, whereas nonvascular moss productivity was especially important in the oldest, most subsided site. The coefficient of variation for soil moisture was higher in the oldest, most subsided site suggesting that in addition to more wet microsites, there were other microsites that were drier. Across all sites, graminoids preferred the cold, dry microsites whereas the moss and shrubs were associated with the warm, moist microsites. Total nitrogen contained in green plant biomass differed across sites, suggesting that there were increases in soil nitrogen availability where permafrost had thawed.     

Warming alters the size spectrum and shifts the distribution of biomass in freshwater ecosystems

Organism size is one of the key determinants of community structure, and its relationship with abundance can describe how biomass is partitioned among the biota within an ecosystem. An outdoor freshwater mesocosm experiment was used to determine how warming of～4 °C would affect the size, biomass and taxonomic structure of planktonic communities. Warming increased the steepness of the community size spectrum by increasing the prevalence of small organisms, primarily within the phytoplankton assemblage and it also reduced the mean and maximum size of phytoplankton by approximately one order of magnitude. The observed shifts in phytoplankton size structure were reflected in changes in phytoplankton community composition, though zooplankton taxonomic composition was unaffected by warming. Furthermore, warming reduced community biomass and total phytoplankton biomass, although zooplankton biomass was unaffected. This resulted in an increase in the zooplankton to phytoplankton biomass ratio in the warmed mesocosms, which could be explained by faster turnover within the phytoplankton assemblages. Overall, warming shifted the distribution of phytoplankton size towards smaller individuals with rapid turnover and low standing biomass, resulting in a reorganization of the biomass structure of the food webs. These results indicate future environmental warming may have profound effects on the structure and functioning of aquatic communities and ecosystems.     

Warming alters community size structure and ecosystem functioning

Global warming can affect all levels of biological complexity, though we currently understand least about its potential impact on communities and ecosystems. At the ecosystem level, warming has the capacity to alter the structure of communities and the rates of key ecosystem processes they mediate. Here we assessed the effects of a 4°C rise in temperature on the size structure and taxonomic composition of benthic communities in aquatic mesocosms, and the rates of detrital decomposition they mediated. Warming had no effect on biodiversity, but altered community size structure in two ways. In spring, warmer systems exhibited steeper size spectra driven by declines in total community biomass and the proportion of large organisms. By contrast, in autumn, warmer systems had shallower size spectra driven by elevated total community biomass and a greater proportion of large organisms. Community-level shifts were mirrored by changes in decomposition rates. Temperature-corrected microbial and macrofaunal decomposition rates reflected the shifts in community structure and were strongly correlated with biomass across mesocosms. Our study demonstrates that the 4°C rise in temperature expected by the end of the century has the potential to alter the structure and functioning of aquatic ecosystems profoundly, as well as the intimate linkages between these levels of ecological organization.     

Temperature alters the relative abundance and population growth rates of species within the Dendroctonus frontalis (Coleoptera: Curculionidae) community

Temperature has strong effects on metabolic processes of individuals and demographics of populations, but effects on ecological communities are not well known. Many economically and ecologically important pest species have obligate associations with other organisms; therefore, effects of temperature on these species might be mediated by strong interactions. The southern pine beetle (Dendroctonus frontalis Zimmermann) harbors a rich community of phoretic mites and fungi that are linked by many strong direct and indirect interactions, providing multiple pathways for temperature to affect the system. We tested the effects of temperature on this community by manipulating communities within naturally infested sections of pine trees. Direct effects of temperature on component species were conspicuous and sometimes predictable based on single-species physiology, but there were also strong indirect effects of temperature via alteration of species interactions that could not have been predicted based on autecological temperature responses. Climatic variation, including directional warming, will likely influence ecological systems through direct physiological effects as well as indirect effects through species interactions.     

Habitat structure and body size distributions: cross‐ecosystem comparison for taxa with determinate and indeterminate growth

Habitat structure across multiple spatial and temporal scales has been proposed as a key driver of body size distributions for associated communities. Thus, understanding the relationship between habitat and body size is fundamental to developing predictions regarding the influence of habitat change on animal communities. Much of the work assessing the relationship between habitat structure and body size distributions has focused on terrestrial taxa with determinate growth, and has primarily analysed discontinuities (gaps) in the distribution of species mean sizes (species size relationships or SSRs). The suitability of this approach for taxa with indeterminate growth has yet to be determined. We provide a cross‐ecosystem comparison of bird (determinate growth) and fish (indeterminate growth) body mass distributions using four independent data sets. We evaluate three size distribution indices: SSRs, species size–density relationships (SSDRs) and individual size–density relationships (ISDRs), and two types of analysis: looking for either discontinuities or abundance patterns and multi‐modality in the distributions. To assess the respective suitability of these three indices and two analytical approaches for understanding habitat–size relationships in different ecosystems, we compare their ability to differentiate bird or fish communities found within contrasting habitat conditions. All three indices of body size distribution are useful for examining the relationship between cross‐scale patterns of habitat structure and size for species with determinate growth, such as birds. In contrast, for species with indeterminate growth such as fish, the relationship between habitat structure and body size may be masked when using mean summary metrics, and thus individual‐level data (ISDRs) are more useful. Furthermore, ISDRs, which have traditionally been used to study aquatic systems, present a potentially useful common currency for comparing body size distributions across terrestrial and aquatic ecosystems.     

Biomass and structure of planktonic communities along an air temperature gradient in subarctic Sweden

1. Air temperature will probably have pronounced effects on the composition of plankton communities in northern lake ecosystems, either via indirect effects on the export of essential elements from catchments or through direct effects of water temperature and the ice‐free period on the behaviour of planktonic organisms. 2. We assessed the role of temperature by comparing planktonic communities in 15 lakes along a 6 °C air temperature gradient in subarctic Sweden. 3. We found that the biomass of phytoplankton, bacterioplankton and the total planktonic biomass were positively related to air temperature, probably as a result of climatic controls on the export of nitrogen from the catchment (which affects phytoplankton biomass) and dissolved organic carbon (affecting bacterioplankton biomass). 4. The structure of the zooplankton community, and top down effects on phytoplankton, were apparently not related to temperature but mainly to trophic interactions ultimately dependent on the presence of fish in the lakes. 5. Our results suggest that air temperature regimes and long‐term warming can have strong effects on the planktonic biomass in high latitude lakes. Effects of temperature on the structure of the planktonic community might be less evident unless warming permits the invasion of fish into previous fishless lakes.     

Community structure of fish in lowland streams differ substantially between subtropical and temperate climates

Fish are important in the structuring of other communities and may have large effects on the functioning of aquatic ecosystems. The structure of fish communities, in turn, seems to differ with climate. We compared the characteristics of fish assemblages in lowland streams located in two contrasting climates (cold-temperate Europe and subtropical South America) by use of published and unpublished data on streams of similar depth, width, and slope (n total?=?91 streams). We also selected a subset of seven comparable little-affected streams in the two contrasting climates: temperate (Denmark, 55°?C57°N, Dk) and subtropical (Uruguay, 30°?C35°S, Uy) and compared the fish community structures in relation to environmental characteristics. We then analysed a series of potential explanatory factors behind the patterns observed, in particular the effect of ambient temperature, by comparing temperature-corrected community metabolism. Significantly higher species richness, higher densities, lower biomass, smaller mean body size, and lower mean weight of fish were observed for the subtropical streams than for the temperate streams, both in the literature review and in the subset of streams. Several characteristics of fish assemblages in streams may be explained by direct and indirect effects of temperature. Accordingly, fish in subtropical systems had a temperature-corrected community metabolism I?m^?2 equal to that of fish in temperate systems, indicating that temperature, besides historical factors, is an important driver of different size structures. Our findings concur with differences previously found in littoral areas of shallow lakes, suggesting that these patterns are not restricted to running waters. Our results elucidate how fish community structure might be affected by increases in temperature triggered by climate warming.