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International audience; Climate warming is considered to be among the most serious of anthropogenic stresses to the environment, because it not only has direct effects on biodiversity, but it also exacerbates the harmful effects of other human-mediated threats. The associated consequences are potentially severe, particularly in terms of threats to species preservation, as well as in the preservation of an array of ecosystem services provided by biodiversity. Among the most affected groups of animals are insects—central components of many ecosystems—for which climate change has pervasive effects from individuals to communities. In this contribution to the scientists' warning series, we summarize the effect of the gradual global surface temperature increase on insects, in terms of physiology, behavior, phenology, distribution, and species interactions, as well as the effect of increased frequency and duration of extreme events such as hot and cold spells, fires, droughts, and floods on these parameters. We warn that, if no action is taken to better understand and reduce the action of climate change on insects, we will drastically reduce our ability to build a sustainable future based on healthy, functional ecosystems. We discuss perspectives on relevant ways to conserve insects in the face of climate change, and we offer several key recommendations on management approaches that can be adopted, on policies that should be pursued, and on the involvement of the general public in the protection effort.

Contains fulltext : 292642.pdf (Publisher’s version ) (Open Access) 10 maart 2023

Oxygen bioavailability is declining in aquatic systems worldwide as a result of climate change and other anthropogenic stressors. For aquatic organisms, the consequences are poorly known but are likely to reflect both direct effects of declining oxygen bioavailability and interactions between oxygen and other stressors, including two—warming and acidification— that have received substantial attention in recent decades and that typically accompany oxygen changes. Drawing on the collected papers in this symposium volume (“An Oxygen Perspective on Climate Change”), we outline the causes and consequences of declining oxygen bioavailability. First, we discuss the scope of natural and predicted anthropogenic changes in aquatic oxygen levels. Although modern organisms are the result of long evolutionary histories during which they were exposed to natural oxygen regimes, anthropogenic change is now exposing them to more extreme conditions and novel combinations of low oxygen with other stressors. Second, we identify behavioral and physiological mechanisms that underlie the interactive effects of oxygen with other stressors, and we assess the range of potential organismal responses to oxygen limitation that occur across levels of biological organization and over multiple timescales. We argue that metabolism and energetics provide a powerful and unifying framework for understanding organism-oxygen interactions. Third, we conclude by out-lining a set of approaches for maximizing the effectiveness of future work, including focusing on long-term experiments using biologically realistic variation in experimental factors and taking truly cross-disciplinary and integrative approaches to understanding and predicting future effects.

Body size is central to ecology at levels ranging from organismal fecundity to the functioning of communities and ecosystems. Understanding temperature‐induced variations in body size is therefore of fundamental and applied interest, yet thermal responses of body size remain poorly understood. Temperature–size (T–S) responses tend to be negative (e.g. smaller body size at maturity when reared under warmer conditions), which has been termed the temperature–size rule (TSR). Explanations emphasize either physiological mechanisms (e.g. limitation of oxygen or other resources and temperature‐dependent resource allocation) or the adaptive value of either a large body size (e.g. to increase fecundity) or a short development time (e.g. in response to increased mortality in warm conditions). Oxygen limitation could act as a proximate factor, but we suggest it more likely constitutes a selective pressure to reduce body size in the warm: risks of oxygen limitation will be reduced as a consequence of evolution eliminating genotypes more prone to oxygen limitation. Thus, T–S responses can be explained by the ‘Ghost of Oxygen‐limitation Past’, whereby the resulting (evolved) T–S responses safeguard sufficient oxygen provisioning under warmer conditions, reflecting the balance between oxygen supply and demands experienced by ancestors. T–S responses vary considerably across species, but some of this variation is predictable. Body‐size reductions with warming are stronger in aquatic taxa than in terrestrial taxa. We discuss whether larger aquatic taxa may especially face greater risks of oxygen limitation as they grow, which may be manifested at the cellular level, the level of the gills and the whole‐organism level. In contrast to aquatic species, terrestrial ectotherms may be less prone to oxygen limitation and prioritize early maturity over large size, likely because overwintering is more challenging, with concomitant stronger end‐of season time constraints. Mechanisms related to time constraints and oxygen limitation are not mutually exclusive explanations for the TSR. Rather, these and other mechanisms may operate in tandem. But their relative importance may vary depending on the ecology and physiology of the species in question, explaining not only the general tendency of negative T–S responses but also variation in T–S responses among animals differing in mode of respiration (e.g. water breathers versus air breathers), genome size, voltinism and thermally associated behaviour (e.g. heliotherms).

Rising temperatures are associated with reduced body size in many marine species, but the biological cause and generality of the phenomenon is debated. We derive a predictive model for body size responses to temperature and oxygen (O 2 ) changes based on thermal and geometric constraints on organismal O 2 supply and demand across the size spectrum. The model reproduces three key aspects of the observed patterns of intergenerational size reductions measured in laboratory warming experiments of diverse aquatic ectotherms (i.e., the “temperature-size rule” [TSR]). First, the interspecific mean and variability of the TSR is predicted from species’ temperature sensitivities of hypoxia tolerance, whose nonlinearity with temperature also explains the second TSR pattern—its amplification as temperatures rise. Third, as body size increases across the tree of life, the impact of growth on O 2 demand declines while its benefit to O 2 supply rises, decreasing the size dependence of hypoxia tolerance and requiring larger animals to contract by a larger fraction to compensate for a thermally driven rise in metabolism. Together our results support O 2 limitation as the mechanism underlying the TSR, and they provide a physiological basis for projecting ectotherm body size responses to climate change from microbes to macrofauna. For small species unable to rapidly migrate or evolve greater hypoxia tolerance, ocean warming and O 2 loss in this century are projected to induce >20% reductions in body mass. Size reductions at higher trophic levels could be even stronger and more variable, compounding the direct impact of human harvesting on size-structured ocean food webs.

The growing use of functional traits in ecological research has brought new insights into biodiversity responses to global environmental change. However, further progress depends on overcoming three major challenges involving (a) statistical correlations between traits, (b) phylogenetic constraints on the combination of traits possessed by any single species, and (c) spatial effects on trait structure and trait–environment relationships. Here, we introduce a new framework for quantifying trait correlations, phylogenetic constraints and spatial variability at large scales by combining openly available species’ trait, occurrence and phylogenetic data with gridded, high‐resolution environmental layers and computational modelling. Our approach is suitable for use among a wide range of taxonomic groups inhabiting terrestrial, marine and freshwater habitats. We demonstrate its application using freshwater macroinvertebrate data from 35 countries in Europe. We identified a subset of available macroinvertebrate traits, corresponding to a life‐history model with axes of resistance, resilience and resource use, as relatively unaffected by correlations and phylogenetic constraints. Trait structure responded more consistently to environmental variation than taxonomic structure, regardless of location. A re‐analysis of existing data on macroinvertebrate communities of European alpine streams supported this conclusion, and demonstrated that occurrence‐based functional diversity indices are highly sensitive to the traits included in their calculation. Overall, our findings suggest that the search for quantitative trait–environment relationships using single traits or simple combinations of multiple traits is unlikely to be productive. Instead, there is a need to embrace the value of conceptual frameworks linking community responses to environmental change via traits which correspond to the axes of life‐history models. Through a novel integration of tools and databases, our flexible framework can address this need.

QUESTIONS: Sod cutting has been used extensively as an effective measure in removing excess N and restoring dwarf shrub dominance in heathlands affected by increased nitrogen deposition. However, recovery of other plant species is often very limited. One barrier is high soil acidity following sod cutting, which results in soil aluminium (Al³⁺) and ammonium (NH₄⁺) reaching toxic concentrations. Sod‐cutting management also removes most of the major nutrients from the system, so intensified nutrient limitation could be an additional barrier to the recovery of species‐rich communities. Soil phosphorus (P) is of special interest as research indicates sod‐cutting management can shift the system to P limitation. LOCATION: Hoge Veluwe National Park, The Netherlands. METHODS: We set up a full‐factorial experiment in sod‐cut heathland formerly encroached by Molinia caerulea, adding phosphate (P+) and lime (Ca+) and over the next three growing seasons, we recorded soil chemistry and plant responses. RESULTS: Soil inorganic N, especially soil NH₄⁺, strongly declined after liming compared to the control situation, confirming that liming alleviates NH₄⁺ toxicity. Addition of P resulted in a similar decline, also suggesting a role for soil phosphate availability in this process. Acid‐sensitive plant species richness increased significantly in both Ca+ and P+ treatment, whereas acid‐insensitive plant species richness only increased significantly as a result of P+ treatment. Mean vegetation relative growth rate increased significantly in both Ca+ and P+ treatments. CONCLUSIONS: Excessive P removal due to sod cutting constitutes a second barrier to restoration in addition to soil acidity. We discourage the large‐scale use of sod cutting to reduce soil N availability in heathlands and propose to use interventions that leave the soil nutrient balance intact, such as burning and grazing, mowing or litter removal, in combination with measures that restore the soil buffer capacity.

There is a growing interest in the physiology underpinning heat tolerance of ectotherms and their responses to the ongoing rise in temperature. However, there is no consensus about the underlying physiological mechanisms. According to “the maintain aerobic scope and regulate oxygen supply” hypothesis, responses to warming at different organizational levels contribute to the ability to safeguard energy metabolism via aerobic pathways. At the cellular level, a decrease in cell size increases the capacity for the uptake of resources (e.g., food and oxygen), but the maintenance of electrochemical gradients across cellular membranes implies greater energetic costs in small cells. In this study, we investigated how different rearing temperatures affected cell size and heat tolerance in the fruit fly Drosophila melanogaster. We tested the hypothesis that smaller‐celled flies are more tolerant to acute, intense heat stress whereas larger‐celled flies are more tolerant to chronic, mild heat stress. We used the thermal tolerance landscape framework, which incorporates the intensity and duration of thermal challenge. Rearing temperatures strongly affected both cell size and survival times. We found different effects of developmental plasticity on tolerance to either chronic or acute heat stress. Warm‐reared flies had both smaller cells and exhibited higher survival times under acute, intense heat stress when compared to cold‐reared flies. However, under chronic, mild heat stress, the situation was reversed and cold‐reared flies, consisting of larger cells, showed better survival. These differences in heat tolerance could have resulted from direct effects of rearing temperature or they may be mediated by the correlated changes in cell size. Notably, our results are consistent with the idea that a smaller cell size may confer tolerance to acute temperatures via enhanced oxygen supply, while a larger cell may confer greater tolerance to chronic and less intense heat stress via more efficient use of resources.

Many ectotherms species grow faster but attain a smaller body size when reared under warmer conditions, a phenomenon known as the Temperature-Size Rule (TSR). This rule appears to be stronger in aquatic ectotherms than in terrestrial ectotherms. The difference could be related to difficulties for oxygen uptake in water, whereas on land, adaptive responses in body size may relate to seasonal time constraints. To assess the role of seasonal time constraints in temperature size response of terrestrial ectotherms, we reared the small copper

Being composed of small cells may carry energetic costs related to maintaining ionic gradients across cell membranes as well as benefits related to diffusive oxygen uptake. Here we test the hypothesis that these costs and benefits of cell size in ectotherms are temperature dependent. To study the consequences of cell size for whole-organism metabolic rate we compared diploid and triploid zebrafish larvae differing in cell size. A fully factorial design was applied combining three different rearing and test temperatures that allowed us to distinguish acute from acclimated thermal effects. Individual oxygen consumption rates of diploid and triploid larvae across declining levels of oxygen availability were measured. We found that both acute and acclimated thermal effects affected the metabolic response. In comparison to triploids, diploids responded more strongly to acute temperatures, especially when reared at the highest temperature. These observations support the hypothesis that animals composed of smaller cells (i.e. diploids) are less vulnerable to oxygen limitation in warm aquatic habitats. Furthermore, we found slightly improved hypoxia tolerance in diploids. By contrast, warm-reared triploids had higher metabolic rates when they were tested at acute cold temperature, suggesting that being composed of larger cells may provide metabolic advantages in the cold. We offer two mechanisms as a potential explanation of this result, related to homeoviscous adaptation of membrane function and the mitigation of developmental noise. Our results suggest that being composed of larger cells provides metabolic advantages in cold water, while being composed of smaller cells provides metabolic advantages in warm water.

Trait-based approaches are promising to make generalizations about the sensitivity of species and populations to pesticides. Two traits that may shape the sensitivity to pesticides are the surface area (related to pesticide uptake) and the metabolic rate (related to pesticide elimination). We compared the sensitivity of damselfly larvae to the pesticide chlorpyrifos and how this was modified by loss of external gills (autotomy, reducing the surface area) in both fast pace-of-life (high metabolic rate) and slow pace-of-life (low metabolic rate) populations of Ischnura elegans. The slow-paced populations were more sensitive to the pesticide than the fast-paced populations in terms of survival, growth and energy metabolism. This suggests the higher metabolic rate of fast-paced populations enabled a faster pesticide elimination. Pesticide exposure also reduced heat tolerance, especially in slow-paced larvae under hypoxia. Gill loss had opposite effects on pesticide sensitivity in slow- and fast-paced populations. In slow-paced larvae, gill loss lowered the sensitivity to the pesticide, while in fast-paced larvae, gill loss increased the sensitivity. This difference likely reflects the balance between the roles of the gills in pesticide uptake (more detrimental in slow-paced populations) and oxygen uptake (more important in fast-paced populations). Our results highlight the need to consider trait interactions when applying trait-based approaches to predict the sensitivity to pesticides. ispartof: ENVIRONMENTAL POLLUTION vol:282 ispartof: location:England status: published

International audience; Diving as a lifestyle has evolved on multiple occasions when air-breathing terrestrial animals invaded the aquatic realm, and diving performance shapes the ecology and behaviour of all air-breathing aquatic taxa, from small insects to great whales. Using the largest dataset yet assembled, we show that maximum dive duration increases predictably with body mass in both ectotherms and endotherms. Compared to endotherms, ectotherms can remain submerged for longer, but the mass scaling relationship for dive duration is much steeper in endotherms than in ectotherms. These differences in diving allometry can be fully explained by inherent differences between the two groups in their metabolic rate and how metabolism scales with body mass and temperature. Therefore, we suggest that similar constraints on oxygen storage and usage have shaped the evolutionary ecology of diving in all air-breathing animals, irrespective of their evolutionary history and metabolic mode. The steeper scaling relationship between body mass and dive duration in endotherms not only helps explain why the largest extant vertebrate divers are endothermic rather than ectothermic, but also fits well with the emerging consensus that large extinct tetrapod divers (e.g. plesiosaurs, ichthyosaurs and mosasaurs) were endothermic.

Temperature has a profound impact on ectotherms. Warming increases the metabolic oxygen demand of ectotherms, which could result in a mismatch between their oxygen demand and their ability to extract and deliver sufficient oxygen to meet demand. This hypothesis has been mainly tested using short-term exposure to intense thermal stress. However, the thermal responses of organisms can be different on longer timescales, where physiological acclimation becomes increasingly important. Such thermal acclimation effects may reduce the vulnerability of ectotherms to warming on the long term. Thus, responses to intense, short-term thermal stress may be different from responses to moderate, prolonged thermal stress. Here, we examine the effect of thermal acclimation on heat tolerance and metabolism in the aquatic ectotherm Gammarus fossarum (Koch, 1836). Amphipods were acclimated to either 11.1 ± 0.1 °C or 19.8 ± 0.1 °C and after thermal acclimation we measured both their metabolism and their survival time at different temperatures. Our results show that metabolism strongly increased with increasing temperatures in the cold-acclimated group, but less so in the warm-acclimated group. Cold-acclimated amphipods were also more sensitive to thermal stress, especially during prolonged exposure. Thus, the differences between both thermal acclimation groups support the idea of oxygen-limited heat tolerance: cold-acclimated amphipods showed increased oxygen consumption and decreased thermal tolerance. However, across individuals, those that sharply increased oxygen consumption with increasing temperature did not differ in heat tolerance from individuals whose metabolism was much less sensitive to temperature. Thus, acclimation to different temperatures appeared to be beneficial, but a role for oxygen limitation could not be demonstrated unambiguously. Beneficial effect of acclimation were much larger during prolonged exposure, with the acclimation response ratio (ARR) ranging from 0.03 to over 0.5 depending on the time scale (minutes to months). Thus, the acclimatory capacity may have been underestimated by short-term experimental studies.

Thermal history can plastically alter the response of ectotherms to temperature, and thermal performance curves (TPCs) are powerful tools for exploring how organismal-level performance varies with temperature. Plasticity in TPCs may be favoured in thermally variable habitats, where it can result in fitness benefits. However, thermal physiology remains insufficiently studied for freshwater insects despite freshwater biodiversity being at great risk under global change. Here, we assess how acclimation at either summer or winter average temperatures changes TPCs for locomotion activity and metabolism in Enochrus jesusarribasi (Hydrophilidae), a water beetle endemic to shallow saline streams in SE Spain. This beetle is a bimodal gas exchanger and so we also assessed how aerial and aquatic gas exchange varied across temperatures for both acclimation treatments. Responses of locomotory TPCs to thermal acclimation were relatively weak, but high temperature acclimated beetles tended to exhibit higher maximum locomotor activity and reduced TPC breadth than those acclimated at lower temperature. High temperature acclimation increased the thermal sensitivity of metabolic rates, contrary to the response generally found in aquatic organisms. Higher metabolic rates upon high temperature acclimation were achieved by increasing aerial, rather than aquatic oxygen uptake. Such plastic respiratory behaviour likely contributed to enhanced locomotor performance at temperatures around the optimum and thermal plasticity could thus be an important component in the response of aquatic insects to climate change. However, high temperature acclimation appeared to be detrimental for locomotion in subsequent exposure at upper sublethal temperatures, suggesting that this narrow range endemic may be vulnerable to future climate warming. This study demonstrates that TPCs are context-specific, differing with performance metric as well as thermal history. Such context dependency must be considered when using TPCs to predict organismal responses to climate change.

Contains fulltext : 193296.pdf (Publisher’s version ) (Open Access) 13 p.

Summary Globalization and climate change trigger species invasions and range shifts, which reshuffle communities at an exceptional rate and expose plant migrants to unfamiliar herbivores. Dominant hypotheses to predict plant success are based on evolutionary novelty: either herbivores are maladapted to consume novel plants (enemy release hypothesis), or novel plants are maladapted to deter herbivores (biotic resistance hypothesis). Since novelty can work both ways, it fails to consistently predict when herbivores will consume novel over non-novel plants. Surprisingly, the value of using plant traits to predict herbivore consumption of novel plants remains largely unexplored. We hypothesized that (i) plant traits explain generalist herbivore consumption rates of novel and non-novel plants, and (ii) any effect of novelty will be grounded in consistent trait differences between native and novel plants. Lastly, we expected to find (iii) differences in plant traits and plant consumption rates across latitude. To test these hypotheses, we measured the consumption rate of plant species for a tropical and a temperate generalist herbivore in controlled feeding trials by offering them a large variety of 40 plant species from different geographical origins. Therefore, whether a plant was novel depended on the herbivore used, allowing us to disentangle plant identity from plant novelty. We also measured plant chemical traits and determined whether traits, geographical origin or novelty best explained herbivore consumption rates. Both generalist herbivores consumed more of plants with a high nitrogen-to-phenolic compounds ratio, irrespective of the plant's novelty to the herbivore. A pattern of increasing plant's nitrogen-to-phenolics ratio with latitude could explain why both the tropical and temperate herbivore consumed more of plants from temperate regions. Plant novelty and its geographical origin no longer explained consumption rates once differences in nitrogen-to-phenolic compounds ratio were taken into account. We show that differences in plant traits along a latitudinal cline determine herbivore consumption rates, irrespective of whether plants are novel or familiar. Therefore, we propose that integrating evolutionary novelty theory with plant traits and biogeography will increase our understanding of the consequences of plant species migration beyond biogeographical barriers. A lay summary is available for this article.

Contains fulltext : 216856.pdf (Publisher’s version ) (Open Access) There is renewed interest in the regulation and consequences of cell size adaptations in studies on understanding the ecophysiology of ectotherms. Here we test if induction of triploidy, which increases cell size in zebrafish (Danio rerio), makes for a good model system to study consequences of cell size. Ideally, diploid and triploid zebrafish should differ in cell size, but should otherwise be comparable in order to be suitable as a model. We induced triploidy by cold shock and compared diploid and triploid zebrafish larvae under standard rearing conditions for differences in genome size, cell size and cell number, development, growth and swimming performance and expression of housekeeping genes and hsp70.1. Triploid zebrafish have larger but fewer cells, and the increase in cell size matched the increase in genome size (+ 50%). Under standard conditions, patterns in gene expression, ontogenetic development and larval growth were near identical between triploids and diploids. However, under demanding conditions (i.e. the maximum swimming velocity during an escape response), triploid larvae performed poorer than their diploid counterparts, especially after repeated stimuli to induce swimming. This result is consistent with the idea that larger cells have less capacity to generate energy, which becomes manifest during repeated physical exertion resulting in increased fatigue. Triploidy induction in zebrafish appears a valid method to increase specifically cell size and this provides a model system to test for consequences of cell size adaptation for the energy budget and swimming performance of this ectothermic vertebrate. 02 maart 2020

Significance Organismal responses to climate change are mediated through its effects on physiology and metabolism. In aquatic environments, both water temperature and oxygen availability may modulate these responses by altering the aerobic metabolism fueling physiological performance. However, ecological models aimed at predicting how environmental factors shape aerobic metabolism disregard the role of oxygen supply. Here, we expand on these models by explicitly incorporating oxygen supply. Our results show that warmer water increases oxygen demand relative to supply, and the resulting reduction in aerobic scope appears to be stronger in larger individuals. Smaller aerobic scopes in warming water imply that climate change will reduce energy budgets needed to support the activities of aquatic animals and their physiological performance in the future., Both oxygen and temperature are fundamental factors determining metabolic performance, fitness, ecological niches, and responses of many aquatic organisms to climate change. Despite the importance of physical and physiological constraints on oxygen supply affecting aerobic metabolism of aquatic ectotherms, ecological theories such as the metabolic theory of ecology have focused on the effects of temperature rather than oxygen. This gap currently impedes mechanistic models from accurately predicting metabolic rates (i.e., oxygen consumption rates) of aquatic organisms and restricts predictions to resting metabolism, which is less affected by oxygen limitation. Here, we expand on models of metabolic scaling by accounting for the role of oxygen availability and temperature on both resting and active metabolic rates. Our model predicts that oxygen limitation is more likely to constrain metabolism in larger, warmer, and active fish. Consequently, active metabolic rates are less responsive to temperature than are resting metabolic rates, and metabolism scales to body size with a smaller exponent whenever temperatures or activity levels are higher. Results from a metaanalysis of fish metabolic rates are consistent with our model predictions. The observed interactive effects of temperature, oxygen availability, and body size predict that global warming will limit the aerobic scope of aquatic ectotherms and may place a greater metabolic burden on larger individuals, impairing their physiological performance in the future. Our model reconciles the metabolic theory with empirical observations of oxygen limitation and provides a formal, quantitative framework for predicting both resting and active metabolic rate and hence aerobic scope of aquatic ectotherms.

Assemblages of aquatic insects are structured by multiple biotic and abiotic conditions, including temperature, salinity and oxygen. Here we highlight recent developments in our understanding of how high temperatures, elevated salinities and low oxygen levels affect physiological processes, responses at the organismal level, and impacts on species interaction and community assembly. As aquatic insects may be exposed to multiple stressors, we review their sensitivity to interactive effects of multiple stressors. While each of these stressors may operate via different physiological mechanisms, they all influence the overall energy budget as well as the allocation of energy to competing functions such as homeostatic maintenance, growth, development and reproduction. As such, there is potential for interaction whereby one stressor may exacerbate the effect of another stressor. Integrating research on these stressors can provide a powerful approach for delineating the sensitivity of aquatic insects to multiple stressors and developing sound management practices.

The ability of organisms to cope with environmental stressors depends on the duration and intensity of the stressor, as well as the type of stress. For aquatic organisms, oxygen limitation has been implicated in limiting heat tolerance. Here we examine how starvation affects heat tolerance in the amphipod Gammarus fossarum (Koch, 1836) and whether observed changes can be explained from alterations in oxidative metabolism, depletion of energy reserves, upregulation of heat shock proteins or susceptibility to oxygen limitation. Starved amphipods showed impaired survival compared to fed amphipods during prolonged exposure to mild heat. In contrast, under acute, high-intensity heat exposure they actually showed improved survival. We observed a lower demand for oxygen in starved amphipods which could make them less susceptible to oxygen limitation. Such a role for oxygen in limiting heat tolerance was verified as hypoxia impaired the heat tolerance of amphipods, especially starved ones. Fed amphipods likely rely more on anaerobic metabolism to maintain energy status during heat stress, whereas for starved amphipods aerobic metabolism appears to be more important. The depletion of their energy reserves constrains their ability to maintain energy status via anaerobic metabolism. We did not find evidence that alterations in heat tolerance following starvation were related to the upregulation of heat shock proteins. In conclusion, starvation can have opposite effects on heat tolerance, acting via pathways that are operating on different time scales.

Global warming appears to favour smaller-bodied organisms, but whether larger species are also more vulnerable to thermal extremes, as suggested for past mass-extinction events, is still an open question. Here, we tested whether interspecific differences in thermal tolerance (heat and cold) of ectotherm organisms are linked to differences in their body mass and genome size (as a proxy for cell size). Since the vulnerability of larger, aquatic taxa to warming has been attributed to the oxygen limitation hypothesis, we also assessed how body mass and genome size modulate thermal tolerance in species with contrasting breathing modes, habitats and life-stages. A database with the upper (CTmax) and lower (CTmin) critical thermal limits and their methodological aspects was assembled comprising more than 500 species of ectotherms. Our results demonstrate that thermal tolerance in ectotherms is dependent on body mass and genome size and these relationships became especially evident in prolonged experimental trials where energy efficiency gains importance. During long-term trials, CTmax was impaired in larger-bodied water-breathers, consistent with a role for oxygen limitation. Variation in CTmin was mostly explained by the combined effects of body mass and genome size and it was enhanced in larger-celled, air-breathing species during long-term trials, consistent with a role for depolarization of cell membranes. Our results highlight the importance of accounting for phylogeny and exposure duration. Especially when considering long-term trials, the observed effects on thermal limits are more in line with the warming-induced reduction in body mass observed during long-term rearing experiments.

Temperature affects the physiology and life-history of ectothermic animals, often increasing metabolic rate and decreasing body size. Oxygen limitation has been put forward as a mechanism to explain thermal responses of body size and the ability to survive stress. However the time-scales involved in growth performance and heat tolerance differ radically. In order to increase our understanding of oxygen and temperature effects on body size and heat tolerance and the time scale involved, we reared Lymnaea stagnalis under six combinations of temperature and oxygen tension from hatching up to an age of 300 days and recorded shell length during this whole period. At the end of this period, we determined scope for growth by measuring food intake rate, assimilation efficiency, respiration rate and ammonium excretion rate at two different temperatures. We also measured the snails’ ability to survive heat stress (CTmax), both at normoxia and hypoxia. We found that scope for growth and long term growth performance were much more affected by interactions of chronic oxygen and temperature conditions during rearing than by acute conditions during testing. Furthermore, our study shows that individual variation in growth performance can be traced back to individual differences in rates of food and oxygen consumption. Developmental acclimation also gave rise to differences in CTmax, but these were relatively small and were only expressed when CTmax was tested under hypoxia. The large effects of rearing oxygen conditions on growth and other physiological rates compared to modest effects of test oxygen conditions on CTmax suggest that small effects of hypoxia on the short term (e.g. heat tolerance) may nevertheless have large repercussions on the long term (e.g. growth and reproduction), even in a pulmonate snail that can compensate for hypoxia to some extent by aerial respiration.

How seed-eating animals influence the germination of endozoochorous plant species is of great relevance to our understanding of long-distance seed dispersal. To evaluate the effect of gut passage on the germination speed of aquatic and riparian plants, we fed seeds of 19 species to common carp ( Cyprinus carpio ) and Mozambique tilapia ( Oreochromis mossambicus ). Carp has a pharyngeal ‘mill’, compensating for lack of a stomach and teeth, while tilapia has tiny teeth and an acid stomach. Potamogeton species, Alisma plantago-aquatica and Sagittaria sagittifolia showed a continuous germination with relatively low, and constant germination rates throughout the germination period. In contrast, the seeds of Myriophyllum spicatum and Nymphoides peltata and those of the riparian species exhibited discontinuous germination, i.e. almost all seeds germinated within a few days. Compared to control seeds, we found on average a negative effect of gut passage on the germination speed for carp, but no effect for tilapia. However, several plant species deviated from this general pattern. For example, gut passage in carp enhanced germination speed of viable seeds relative to controls for Potamogeton alpinus , but reduced it for Filipendula ulmaria . Similarly, tilapia enhanced germination speed for P . alpinus , but reduced it for P. pusillus . We conclude that carp’s efficient mastication affects the germination speed of plant species after gut passage, whereas stomach-passage in tilapia generally does not influence the germination speed when compared to control seeds. However, inherent variation across species in germination speed proved to be greater than effects of fish gut passage.

Linking variation in species' traits to large-scale environmental gradients can lend insight into the evolutionary processes that have shaped functional diversity and future responses to environmental change. Here, we ask how heat and cold tolerance vary as a function of latitude, elevation and climate extremes, using an extensive global dataset of ectotherm and endotherm thermal tolerance limits, while accounting for methodological variation in acclimation temperature, ramping rate and duration of exposure among studies. We show that previously reported relationships between thermal limits and latitude in ectotherms are robust to variation in methods. Heat tolerance of terrestrial ectotherms declined marginally towards higher latitudes and did not vary with elevation, whereas heat tolerance of freshwater and marine ectotherms declined more steeply with latitude. By contrast, cold tolerance limits declined steeply with latitude in marine, intertidal, freshwater and terrestrial ectotherms, and towards higher elevations on land. In all realms, both upper and lower thermal tolerance limits increased with extreme daily temperature, suggesting that different experienced climate extremes across realms explain the patterns, as predicted under the Climate Extremes Hypothesis . Statistically accounting for methodological variation in acclimation temperature, ramping rate and exposure duration improved model fits, and increased slopes with extreme ambient temperature. Our results suggest that fundamentally different patterns of thermal limits found among the earth's realms may be largely explained by differences in episodic thermal extremes among realms, updating global macrophysiological ‘rules’. This article is part of the theme issue ‘Physiological diversity, biodiversity patterns and global climate change: testing key hypotheses involving temperature and oxygen’.

For aquatic breathers, hypoxia and warming can act synergistically causing a mismatch between oxygen supply (reduced by hypoxia) and oxygen demand (increased by warming). The vulnerability of these species to such interactive effects may differ during ontogeny due to differing gas exchange systems. This study examines respiratory responses to temperature and hypoxia across four life-stages of the intertidal porcelain crab Petrolisthes laevigatus. Eggs, megalopae, juveniles and adults were exposed to combinations of temperatures from 6 to 18 °C and oxygen tensions from 2 to 21 kPa. Metabolic rates differed strongly across life-stages which could be partly attributed to differences in body mass. However, eggs exhibited significantly lower metabolic rates than predicted for their body mass. For the other three stages, metabolic rates scaled with a mass exponent of 0.89. Mass scaling exponents were similar across all temperatures, but were significantly influenced by oxygen tension (the highest at 9 and 14 kPa, and the lowest at 2 kPa). Respiratory responses across gradients of oxygen tension were used to calculate the response to hypoxia, whereby eggs, megalopae and juveniles responded as oxyconformers and adults as oxyregulators. The thermal sensitivity of the metabolic rates (Q10) were dependent on the oxygen tension in megalopae, and also on the interaction between oxygen tension and temperature intervals in adults. Our results thus provide evidence on how the oxygen tension can modulate the mass dependence of metabolic rates and demonstrate changes in respiratory control from eggs to adults. In light of our results indicating that adults show a good capacity for maintaining metabolism independent of oxygen tension, our study highlights the importance of assessing responses to multiple stressors across different life-stages to determine how vulnerability to warming and hypoxia changes during development. Electronic supplementary material The online version of this article (10.1007/s00227-018-3406-z) contains supplementary material, which is available to authorized users.

Predicting species responses to global warming is the holy grail of climate change science. As temperature directly affects physiological rates, it is clear that a mechanistic understanding of species vulnerability should be grounded in organismal physiology. Here, we review what respiratory physiology can offer the field of thermal ecology, showcasing different perspectives on how respiratory physiology can help explain thermal niches. In water, maintaining adequate oxygen delivery to fuel the higher metabolic rates under warming conditions can become the weakest link, setting thermal tolerance limits. This has repercussions for growth and scaling of metabolic rate. On land, water loss is more likely to become problematic as long as O2 delivery and pH balance can be maintained, potentially constraining species in their normal activity. Therefore, high temperatures need not be lethal, but can still affect the energy intake of an animal, with concomitant consequences for long-term fitness. While respiratory challenges and adaptive responses are diverse, there are clear recurring elements such as oxygen uptake, CO2 excretion, and water homeostasis. We show that respiratory physiology has much to offer the field of thermal ecology and call for an integrative, multivariate view incorporating respiratory challenges, thermal responses, and energetic consequences. Fruitful areas for future research are highlighted.

Acidification has led to a strong decline of species characteristic of shallow soft-water lakes. In spite of reductions in acidifying deposition, natural recovery of biodiversity is modest or even absent, suggesting that the impact of acidification is difficult to reverse. We compared recovery from acidification in non-restored and restored lakes using data from 1983 and 2004. In restored lakes, accumulated organic matter was removed and alkaline water was supplied, resulting in an increase in pH and alkalinity and a decrease in ammonium, sulphur and aluminium. For evaluation of biotic changes we selected chironomid larvae (Diptera). In restored lakes, rarefied species richness increased, chironomid species composition changed and responses of chironomid taxa and their life-history strategies indicated a shift towards pre-acidification assemblages. Species adapted to dynamic and stressful environments decreased in favour of those with life history strategies suitable for more benign environments. In non-restored lakes, chironomid response did not indicate a recovery, despite an improved water chemistry in terms of decreased acidity and sulphur (not ammonium and aluminium). Instead, stressful conditions related to oxygen shortage became more prevalent as a decrease was observed in the chironomids least adapted to periods with low oxygen availability. Acidification has inhibited decomposition, resulting in the accumulation of organic material. Natural recovery from acidification resumed and increased the decomposition of this accumulated organic material, resulting in release of nutrients, consumption of oxygen and a decline of sensitive bottom-dwelling fauna such as chironomids. Therefore, active restoration by removal of accumulated organic matter from sand bottoms is essential for a recovery of chironomid assemblages.

In the face of ongoing habitat fragmentation, species–area relationships (SARs) have gained renewed interest and are increasingly used to set conservation priorities. An important question is how large habitat areas need to be to optimize biodiversity conservation. The relationship between area and species richness is explained by colonization–extinction dynamics, whereby smaller sites harbor smaller populations, which are more prone to extinction than the larger populations sustained by larger sites. These colonization–extinction dynamics are predicted to vary with trophic rank, habitat affinity, and dispersal ability of the species. However, empirical evidence for the effect of these species characteristics on SARs remains inconclusive.In this study we used carabid beetle data from 58 calcareous grassland sites to investigate how calcareous grassland area affects species richness and activity density for species differing in trophic rank, habitat affinity, and dispersal ability. In addition, we investigated how SARs are affected by the availability of additional calcareous grassland in the surrounding landscape.Beetle species richness and activity density increased with calcareous grassland area for zoophagous species that are specialists for dry grasslands and, to a lesser extent, for zoophagous habitat generalists. Phytophagous species and zoophagous forest and wet-grassland specialists were not affected by calcareous grassland area. The dependence of species on large single sites increased with decreasing dispersal ability for species already vulnerable to calcareous grassland area. Additional calcareous grassland in the landscape had a positive effect on local species richness of both dry-grassland specialists and generalists, but this effect was restricted to a few hundred meters.Our results demonstrate that SARs are affected by trophic rank, habitat affinity, and dispersal ability. These species characteristics do not operate independently, but should be viewed in concert. In addition, species' responses depend on the landscape context. Our study suggests that the impact of habitat area on trophic interactions may be larger than previously anticipated. In small habitat fragments surrounded by a hostile matrix, food chains may be strongly disrupted. This highlights the need to conserve continuous calcareous grassland patches of at least several hectares in size.Read More: http://www.esajournals.org/doi/abs/10.1890/14-0082.1

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Graphical abstract, Highlights • Heat tolerance and its plasticity are positively correlated in Deronectes beetles. • Deronectes beetles have setal tracheal gills which support under water respiration. • Gill density may explain differences in thermal tolerance and habitat use. • Habitat conditions may have simultaneously selected for cold tolerance and gills. • Gill density may underlie the relation between heat tolerance and its plasticity., In the face of global warming, both the absolute thermal tolerance of an ectotherm, and its ability to shift its tolerance level via acclimation, are thought to be fundamentally important. Understanding the links between tolerance and its plasticity is therefore critical to accurately predict vulnerability to warming. Previous studies in a number of ectotherm taxa suggest trade-offs in the evolution of thermal tolerance and its plasticity, something which does not, however, apply to Deronectes diving beetles, where these traits are instead positively correlated. Here we revisit the relationship between thermal tolerance and plasticity in these beetles, paying attention to a recently discovered morphological adaptation supporting under water respiration – setal tracheal gills. Hollow setae on the elytra interconnect with the beetle’s tracheal system, providing a gas exchange surface that allows oxygen to be extracted directly from the water. This enables individuals to stay submerged for longer than their subelytral air stores would allow. We show that hypoxia reduced heat tolerance, especially when individuals were denied access to air, forcing them to rely solely on aquatic gas exchange. Species with higher densities of these gas-exchanging setae exhibited improved cold tolerance, but reduced heat tolerance and lower plasticity of heat tolerance. Differences in setal tracheal gill density across species were also related to habitat use: species with low gill density were found mainly in intermittent, warmer rivers, where underwater gas exchange is more problematic and risks of surfacing may be smaller. Moreover, when controlling for differences in gill density we no longer found a significant relationship between heat tolerance and its plasticity, suggesting that the previously reported positive relationship between these variables may be driven by differences in gill density. Differences in environmental conditions between the preferred habitats could simultaneously select for characteristic differences in both thermal tolerance and gill density. Such simultaneous selection may have resulted in a non-causal association between cold tolerance and gill density. For heat tolerance, the correlations with gill density could reflect a causal relationship. Species relying strongly on diffusive oxygen uptake via setal tracheal gills may have a reduced oxygen supply capacity and may be left with fewer options for matching oxygen uptake to oxygen demand during acclimation, which could explain their reduced heat tolerance and limited plasticity. Our study helps shed light on the mechanisms that underpin thermal tolerance and plasticity in diving air-breathing ectotherms, and explores how differences in thermal tolerance across species are linked to their selected habitat, morphological adaptations and evolutionary history.

Insects can experience functional hypoxia, a situation in which O2 supply is inadequate to meet oxygen demand. Assessing when functional hypoxia occurs is complex, because responses are graded, age and tissue dependent, and compensatory. Here, we compare information gained from metabolomics and transcriptional approaches and by manipulation of the partial pressure of oxygen. Functional hypoxia produces graded damage, including damaged macromolecules and inflammation. Insects respond by compensatory physiological and morphological changes in the tracheal system, metabolic reorganization, and suppression of activity, feeding, and growth. There is evidence for functional hypoxia in eggs, near the end of juvenile instars, and during molting. Functional hypoxia is more likely in species with lower O2 availability or transport capacities and when O2 need is great. Functional hypoxia occurs normally during insect development and is a factor in mediating life-history trade-offs.

Pollution and predation are two omnipresent stressors in aquatic systems that can interact in multiple ways, thereby challenging accurate assessment of the effects of pollutants in natural systems. Despite the widespread occurrence of morphological antipredator mechanisms, no studies have tested how these can affect the sensitivity of prey to pesticides. Sensitivity to pesticides is typically measured via reductions in growth rates and survival, but also reductions in heat tolerance are to be expected and are becoming increasingly important in a warming world. We investigated how autotomy, a widespread morphological antipredator mechanism where animals sacrifice a body part (here the caudal lamellae) to escape when attacked by a predator, modified the sensitivity to the insecticide chlorpyrifos in larvae of the damselfly Coenagrion puella. Exposure to chlorpyrifos reduced the growth rate and heat tolerance (measured as CTmax). A key finding was that the pesticide had a greater impact on growth rates of intact animals, i.e. those that retained their lamellae. This reduced sensitivity to chlorpyrifos in animals without lamellae can be explained by the reduced outer surface area which is expected to result in a lower uptake of the pesticide. Larvae that underwent autotomy exhibited a lower heat tolerance, which may also be explained by the reduced surface area and the associated reduction in oxygen uptake. There is a wide diversity of morphological antipredator mechanisms, suggesting that there will be more examples where these mechanisms affect the vulnerability to pollutants. Given the importance of pollution and predation as structuring forces in aquatic food webs, exploring the potential interactions between morphological antipredator mechanisms and sensitivity to pollutants will be crucial for risk assessment of pollutants in aquatic systems. ispartof: Science of the Total Environment vol:626 pages:1230-1235 ispartof: location:Netherlands status: published

Ectotherms tend to grow faster, but reach a smaller size when reared under warmer conditions. This temperature-size rule (TSR) is a widespread phenomenon. Despite the generality of this pattern, no general explanation has been found. We therefore tested the relative importance of two proposed mechanisms for the TSR: (1) a stronger increase in development rate relative to growth rate at higher temperatures, which would cause a smaller size at maturity, and (2) resource limitation placing stronger constraints on growth in large individuals at higher temperatures, which would cause problems with attaining a large size in warm conditions. We raised Daphnia magna at eight temperatures to assess their size at maturity, asymptotic size, and size of their offspring. We used three clonal lines that differed in asymptotic size and growth rate. A resource allocation model was developed and fitted to our empirical data to explore the effect of both mechanisms for the TSR. The genetic lines of D. magna showed different temperature dependence of growth and development rates resulting in different responses for size at maturity. Also, at warm temperatures, growth was constrained in large, but not in small individuals. The resource allocation model could fit these empirical data well. Based on our empirical results and model explorations, the TSR of D. magna at maturity is best explained by a stronger increase in development rate relative to growth rate at high temperature, and the TSR at asymptotic size is best explained by a size-dependent and temperature-dependent constraint on growth, although resource limitation could also affect size at maturity. In conclusion, the TSR can take different forms for offspring size, size at maturity, and asymptotic size and each form can arise from its own mechanism, which could be an essential step toward finding a solution to this century-old puzzle.

In order to predict which species can successfully cope with global warming and how other environmental stressors modulate their vulnerability to climate-related environmental factors, an understanding of the ecophysiology underpinning thermal limits is essential for both conservation biology and invasion biology.Heat tolerance and the extent to which heat tolerance differed with oxygen availability were examined for four native and four alien freshwater peracarid crustacean species, with differences in habitat use across species. Three hypotheses were tested: (1) Heat and lack of oxygen synergistically reduce survival of species; (2) patterns in heat tolerance and the modulation thereof by oxygen differ between alien and native species and between species with different habitat use; (3) small animals can better tolerate heat than large animals, and this difference is more pronounced under hypoxia.To assess heat tolerances under different oxygen levels, animal survival was monitored in experimental chambers in which the water temperature was ramped up (0.25°C min-1). Heat tolerance (CTmax) was scored as the cessation of all pleopod movement, and heating trials were performed under hypoxia (5 kPa oxygen), normoxia (20 kPa) and hyperoxia (60 kPa).Heat tolerance differed across species as did the extent by which heat tolerance was affected by oxygen conditions. Heat-tolerant species, for example, Asellus aquaticus and Crangonyx pseudogracilis, showed little response to oxygen conditions in their CTmax, whereas the CTmax of heat-sensitive species, for example, Dikerogammarus villosus and Gammarus fossarum, was more plastic, being increased by hyperoxia and reduced by hypoxia.In contrast to other studies on crustaceans, alien species were not more heat-tolerant than native species. Instead, differences in heat tolerance were best explained by habitat use, with species from standing waters being heat tolerant and species from running waters being heat sensitive. In addition, larger animals displayed lower critical maximum temperature, but only under hypoxia. An analysis of data available in the literature on metabolic responses of the study species to temperature and oxygen conditions suggests that oxygen conformers and species whose oxygen demand rapidly increases with temperature (low activation energy) may be more heat sensitive.The alien species D. villosus appeared most susceptible to hypoxia and heat stress. This may explain why this species is very successful in colonizing new areas in littoral zones with rocky substrate which are well aerated due to continuous wave action generated by passing ships or prevailing winds. This species is less capable of spreading to other waters which are poorly oxygenated and where C. pseudogracilis is the more likely dominant alien species. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.13050/suppinfo is available for this article.

Food webs are an integral part of every ecosystem on the planet, yet understanding the mechanisms shaping these complex networks remains a major challenge. Recently, several studies suggested that non-trophic species interactions such as habitat modification and mutual isms can be important determinants of food web structure. However, it remains unclear whether these findings generalize across ecosystems, and whether non-trophic interactions affect food webs randomly, or affect specific trophic levels or functional groups. Here, we combine analyses of 58 food webs from seven terrestrial, freshwater and coastal systems to test (1) the general hypothesis that non-trophic facilitation by habitat-forming foundation species enhances food web complexity, and (2) whether these enhancements have either random or targeted effects on particular trophic levels, functional groups, and linkages throughout the food web. Our empirical results demonstrate that foundation species consistently enhance food web complexity in all seven ecosystems. Further analyses reveal that 15 out of 19 food web properties can be well-approximated by assuming that foundation species randomly facilitate species throughout the trophic network. However, basal species are less strongly, and carnivores are more strongly facilitated in foundation species' food webs than predicted based on random facilitation, resulting in a higher mean trophic level and a longer average chain length. Overall, we conclude that foundation species strongly enhance food web complexity through non-trophic facilitation of species across the entire trophic network. We therefore suggest that the structure and stability of food webs often depends critically on non-trophic facilitation by foundation species.

For aquatic and riparian plants, the important role of fish in seed dispersal is increasingly recognised. While the propensity of seeds to disperse is known to be a function of morphological, physical and chemical traits of the seed, in the case of fish-mediated seed dispersal (ichthyochory), it is largely unknown how seed traits modulate the potential for seed ingestion and their subsequent survival through the gut. Furthermore, which seed traits are important may vary among fish species. To evaluate the role of both seed and fish traits in ichthyochory, we fed seeds of 19 aquatic and riparian plant species to fish with differing feeding mechanisms. Cyprinus carpio (common carp) has a pharyngeal ‘mill’, which it uses physically to crush hard food, while Oreochromis mossambicus (Mozambique tilapia) has only tiny oral and pharyngeal teeth and instead relies more on chemical digestion. A number of seed traits, including hardness, size and shape, were important determinants of the potential of seeds for ichthyochory. Certain traits (e.g. seed dimensions) were more important during ingestion, whereas other traits were more important for seed survival and subsequent germination (e.g. seed hardness, mucilaginous coat). Compared to controls, germination of retrieved seeds in carp was lower in 10 and higher in two plant species, whereas for tilapia, it was lower in seven and higher in three species. Overlap between these plant species was low, indicating clear difference between the fish studied in their potential for seed dispersal. Carp increased in size during the experiment and concomitant decreases in seed survival and retrieval were found, suggesting that body size and the correlated bite force is an important fish trait in ichthyochory. Overall, seed hardness, size and shape appear crucial for the survival of seeds passing through the guts of carp and tilapia. Beyond this general pattern, a greater complexity of trait-performance relationships appeared: different seed traits are involved during each of the stages of ichthyochory. Moreover, the importance of seed traits differed between carp and tilapia, with some traits having interactive and contrasting effects in both fish species. Aquatic plants with floating seeds adapted to hydrochorous dispersal were less likely to be dispersed by tilapia than plants with non-floating seeds, suggesting a dispersal trade-off between ichthyochory and hydrochory. Thus, depending on their seed characteristics, fish may offer an additional dispersal route to aquatic and riparian plants.

Ectotherms tend to grow faster, but reach a smaller size when reared under warmer conditions. This temperature‐size rule (TSR) is a widespread phenomenon. Despite the generality of this pattern, no general explanation has been found. We therefore tested the relative importance of two proposed mechanisms for the TSR: (1) a stronger increase in development rate relative to growth rate at higher temperatures, which would cause a smaller size at maturity, and (2) resource limitation placing stronger constraints on growth in large individuals at higher temperatures, which would cause problems with attaining a large size in warm conditions. We raised Daphnia magna at eight temperatures to assess their size at maturity, asymptotic size, and size of their offspring. We used three clonal lines that differed in asymptotic size and growth rate. A resource allocation model was developed and fitted to our empirical data to explore the effect of both mechanisms for the TSR. The genetic lines of D. magna showed different temperature dependence of growth and development rates resulting in different responses for size at maturity. Also, at warm temperatures, growth was constrained in large, but not in small individuals. The resource allocation model could fit these empirical data well. Based on our empirical results and model explorations, the TSR of D. magna at maturity is best explained by a stronger increase in development rate relative to growth rate at high temperature, and the TSR at asymptotic size is best explained by a size‐dependent and temperature‐dependent constraint on growth, although resource limitation could also affect size at maturity. In conclusion, the TSR can take different forms for offspring size, size at maturity, and asymptotic size and each form can arise from its own mechanism, which could be an essential step toward finding a solution to this century‐old puzzle.

Increased atmospheric deposition rates of nitrogen (N) and sulphur (S) are known to affect soil biogeochemistry and cause a decline in plant biodiversity of heathlands. Concomitant declines of heathland invertebrates are mainly attributed to changes in vegetation composition and altered habitat structure. While there may also be effects on animals through altered plant chemistry, these have received little attention up to now. Here, we remedy this by quantifying soil nutrient and acid buffering status, vegetation composition and structure, plant nutrient stoichiometry, and densities and species richness of Diptera and Carabidae in two large heathland systems. Soil acid buffering status appeared to be a key driver for plant P availability. Sod-cutting was found to further increase plant N:P ratios, suggesting increased P-limitation. Vegetation N:P ratio was negatively linked to invertebrate density and species richness, and was found to impact fauna more strongly than vegetation structure and plant species richness. The relationship between invertebrates and plant C:N ratio was weaker and less consistent, suggesting that for invertebrates, plant P is generally more limiting than N. Our results imply that the role of plant stoichiometry is underestimated in explaining declines of heathland invertebrates, and we here provide a novel mechanistic model including this pathway. Management should therefore not only focus on restoring habitat structural complexity, attention should be paid to restoring plant stoichiometry. This can be achieved through restoring biogeochemical soil conditions, especially by mitigating soil acidification, while measures solely focusing on removal of accumulated N by means of sod-cutting should be avoided.

The collapse of Atlantic salmon (Salmo salar) stocks throughout North-Western Europe is generally ascribed to large-scale river regulation, water pollution and over-fishing in the 19th and 20th century. However, other causes have rarely been quantified, especially those acting before the 19th century. By analysing historical fishery, market and tax statistics, independently confirmed by archaeozoological records, we demonstrate that populations declined by up to 90% during the transitional period between the Early Middle Ages (c. 450–900 AD) and Early Modern Times (c. 1600 AD). These dramatic declines coincided with improvements in watermill technology and their geographical expansion across Europe. Our extrapolations suggest that historical Atlantic salmon runs must have once been very abundant indeed. The historical perspective presented here contributes to a better understanding of the primary factors that led to major declines in salmon populations. Such understanding provides an essential basis for the effective ecological rehabilitation of freshwater ecosystems.

Insufficient oxygen delivery to tissues is hypothesised to limit thermal tolerance, but evidence in ectotherms is mixed. We assessed heat tolerance under hypoxia, normoxia and hyperoxia to test whether the extent in which oxygen can lower or increase heat tolerance differed with mode of respiration, comparing gill-breathing caenogastropods and lung-breathing pulmonates with or without an accessory gill. Hypoxia lowered heat tolerance in three of the four pulmonates (Physa fontinalis, Physa acuta and Planorbis carinatus) by 1.2–2.1°C. Hyperoxia, however, did not increase the heat tolerance in any of the pulmonate species. Thus, heat tolerance limits of these pulmonates does not appear to be oxygen limited under normoxia, possibly because of their high capacity to regulate oxygen consumption associated with aerial gas exchange. Instead, other processes may become limiting at thermal extremes such as loss of protein function, loss of membrane stability or neuronal dysfunction. The caenogastropod species tested (Potamopyrgus antipodarum, Bithynia tentaculata) closed their operculum during the warming experiments. This behavioural response prevented us from obtaining clear results. Nevertheless, our results suggested hyperoxia may increase heat tolerance in B. tentaculata. This could be related to its lower capacity to regulate oxygen, owing to its fully aquatic gas exchange mechanism.

Summary Biological traits of organisms are expected to provide increased mechanistic understanding of species–environment relationships. Linking traits to environmental conditions is, however, not straightforward, as traits are interconnected within species and can affect the adaptive value of each other. The aim of our study was to evaluate the importance of these trait interrelationships for understanding environmental responses of freshwater macroinvertebrates. To this end, we investigated whether environmental responses of macroinvertebrates sharing a given trait were consistent or differed according to their taxonomy or to their other traits. We divided the macroinvertebrates into groups based on single traits (49 single-trait modalities), on taxonomy (10 orders) and on their overall trait profile (10 trait profile groups [TPGs], defined using self-organising maps clustering). Abundances of each of these 69 groups were related to 24 environmental variables using boosted regression tree (BRT) modelling, to assess the environmental responses of single traits, orders and TPGs. Cross-validated predictive power (R2) of the BRT models ranged from < 1% to 38%. Environmental responses of macroinvertebrates sharing a given trait were inconsistent and varied according to order and/or TPG. Single-trait responses often reflected the responses of the most abundant taxonomic group expressing the trait, suggesting that analysis of trait responses simply revealed patterns in habitat use by the most abundant species and not necessarily mechanistic relationships. Further, taxa from the same TPG (hence showing large overlap in their traits) but belonging to different orders showed different environmental responses. This indicates that the order a taxon belongs to confers unique information related to its evolutionary history that was not captured by our 49 trait modalities. However, groupings by orders cannot replace trait-based approaches, since TPGs also revealed differences in trait profiles within some orders, which were associated with different environmental responses. Our results highlight the importance of considering multiple rather than single traits when linking macroinvertebrates to environmental variables, including the potential information conveyed by evolutionary history.

Thermal tolerance has been hypothesized to result from a mismatch between oxygen supply and demand. However, the generality of this hypothesis has been challenged by studies on various animal groups, including air-breathing adult insects. Recently, comparisons across taxa have suggested that differences in gas exchange mechanisms could reconcile the discrepancies found in previous studies. Here, we test this suggestion by comparing the behaviour of related insect taxa with different gas exchange mechanisms, with and without access to air. We demonstrate oxygen-limited thermal tolerance in air-breathing adults of the plastron-exchanging water bug Aphelocheirus aestivalis. Ilyocoris cimicoides, a related, bimodal gas exchanger, did not exhibit such oxygen-limited thermal tolerance and relied increasingly on aerial gas exchange with warming. Intriguingly, however, when denied access to air, oxygen-limited thermal tolerance could also be induced in this species. Patterns in oxygen-limited thermal tolerance were found to be consistent across life-history stages in these insects, with nymphs employing the same gas exchange mechanisms as adults. These results advance our understanding of oxygen limitation at high temperatures; differences in the degree of respiratory control appear to modulate the importance of oxygen in setting tolerance limits., Summary: Oxygen does not appear to universally limit thermal tolerance, but instead, oxygen-limited thermal tolerance is context dependent, being related to a species’ capacity to regulate oxygen consumption.

Aquatic ecological responses to climatic warming are complicated by interactions between thermal effects and other environmental stressors such as organic pollution and hypoxia. Laboratory experiments have demonstrated how oxygen limitation can set heat tolerance for some aquatic ectotherms, but only at unrealistic lethal temperatures and without field data to assess whether oxygen shortages might also underlie sublethal warming effects. Here, we test whether oxygen availability affects both lethal and nonlethal impacts of warming on two widespread Eurasian mayflies, Ephemera danica, Müller 1764 and Serratella ignita (Poda 1761). Mayfly nymphs are often a dominant component of the invertebrate assemblage in streams, and play a vital role in aquatic and riparian food webs. In the laboratory, lethal impacts of warming were assessed under three oxygen conditions. In the field, effects of oxygen availability on nonlethal impacts of warming were assessed from mayfly occurrence in 42 293 UK stream samples where water temperature and biochemical oxygen demand were measured. Oxygen limitation affected both lethal and sublethal impacts of warming in each species. Hypoxia lowered lethal limits by 5.5 °C (±2.13) and 8.2 °C (±0.62) for E. danica and S. ignita respectively. Field data confirmed the importance of oxygen limitation in warmer waters; poor oxygenation drastically reduced site occupancy, and reductions were especially pronounced under warm water conditions. Consequently, poor oxygenation lowered optimal stream temperatures for both species. The broad concordance shown here between laboratory results and extensive field data suggests that oxygen limitation not only impairs survival at thermal extremes but also restricts species abundance in the field at temperatures well below upper lethal limits. Stream oxygenation could thus control the vulnerability of aquatic ectotherms to global warming. Improving water oxygenation and reducing pollution can provide key facets of climate change adaptation for running waters.