Global climate change in large European rivers: long‐term effects on macroinvertebrate communities and potential local confounding factors

Aquatic species living in running waters are widely acknowledged to be vulnerable to climate‐induced, thermal and hydrological fluctuations. Climate changes can interact with other environmental changes to determine structural and functional attributes of communities. Although such complex interactions are most likely to occur in a multiple‐stressor context as frequently encountered in large rivers, they have received little attention in such ecosystems. In this study, we aimed at specifically addressing the issue of relative long‐term effects of global and local changes on benthic macroinvertebrate communities in multistressed large rivers. We assessed effects of hydroclimatic vs. water quality factors on invertebrate community structure and composition over 30 years (1979–2008) in the Middle Loire River, France. As observed in other large European rivers, water warming over the three decades (+0.9 °C between 1979–1988 and 1999–2008) and to a lesser extent discharge reduction (?80 m³ s^?1) were significantly involved in the disappearance or decrease in taxa typical from fast running, cold waters (e.g. Chloroperlidae and Potamanthidae). They explained also a major part of the appearance and increase of taxa typical from slow flowing or standing waters and warmer temperatures, including invasive species (e.g. Corbicula sp. and Atyaephyra desmarestii). However, this shift towards a generalist and pollution tolerant assemblage was partially confounded by local improvement in water quality (i.e. phosphate input reduction by about two thirds and eutrophication limitation by almost one half), explaining a significant part of the settlement of new pollution‐sensitive taxa (e.g. the caddisfly Brachycentridae and Philopotamidae families) during the last years of the study period. The regain in such taxa allowed maintaining a certain level of specialization in the invertebrate community despite climate change effects.     

Landscape variables influence taxonomic and trait composition of insect assemblages in Neotropical savanna streams

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Trends in water quality and discharge confound long-term warming effects on river macroinvertebrates

1. Climate‐change effects on rivers and streams might interact with other pressures, such as pollution, but long‐term investigations are scarce. We assessed trends among macroinvertebrates in 50 southern English streams in relation to temperature, discharge and water quality over 18 years (1989–2007). 2. Long‐term records, coupled with estimates from inter‐site calibrations of 3–4 years, showed that mean stream temperatures in the study area had increased by 2.1–2.9 °C in winter and 1.1–1.5 °C in summer over the 26 year period from 1980 to 2006, with trends in winter strongest. 3. While invertebrate assemblages in surface‐fed streams were constant, those in chalk‐streams changed significantly during 1989–2007. Invertebrate trends correlated significantly with temperature, but effects were spurious because (i) assemblages gained taxa typical of faster flow or well‐oxygenated conditions, contrary to expectations from warming; (ii) more invertebrate families increased in abundance than declined and (iii) concomitant changes in water quality (e.g. declining orthophosphate, ammonia and biochemical oxygen demand), or at some sites changes in discharge, explained more variation in invertebrate abundance and composition than did temperature. 4. These patterns were reconfirmed in both group‐ and site‐specific analyses. 5. We conclude that recent winter‐biased warming in southern English chalk‐streams has been insufficient to affect invertebrates negatively over a period of improving water quality. This implies that positive management can minimize some climate‐change impacts on stream ecosystems. Chalk‐stream invertebrates are sensitive, nevertheless, to variations in discharge, and detectable changes could occur if climate change alters flow pattern. 6. Because climatic trends now characterize many inter‐annual time‐series, we caution other investigators to examine whether putative effects on ecological systems are real or linked spuriously to other causes of change.     

Hydrological and environmental variables outperform spatial factors in structuring species,trait composition,and beta diversity of pelagic algae

There has been increasing interest in algae‐based bioassessment, particularly, trait‐based approaches are increasingly suggested. However, the main drivers, especially the contribution of hydrological variables, of species composition, trait composition, and beta diversity of algae communities are less studied. To link species and trait composition to multiple factors (i.e., hydrological variables, local environmental variables, and spatial factors) that potentially control species occurrence/abundance and to determine their relative roles in shaping species composition, trait composition, and beta diversities of pelagic algae communities, samples were collected from a German lowland catchment, where a well‐proven ecohydrological modeling enabled to predict long‐term discharges at each sampling site. Both trait and species composition showed significant correlations with hydrological, environmental, and spatial variables, and variation partitioning revealed that the hydrological and local environmental variables outperformed spatial variables. A higher variation of trait composition (57.0%) than species composition (37.5%) could be explained by abiotic factors. Mantel tests showed that both species and trait‐based beta diversities were mostly related to hydrological and environmental heterogeneity with hydrological contributing more than environmental variables, while purely spatial impact was less important. Our findings revealed the relative importance of hydrological variables in shaping pelagic algae community and their spatial patterns of beta diversities, emphasizing the need to include hydrological variables in long‐term biomonitoring campaigns and biodiversity conservation or restoration. A key implication for biodiversity conservation was that maintaining the instream flow regime and keeping various habitats among rivers are of vital importance. However, further investigations at multispatial and temporal scales are greatly needed.     

Impacts of climatic stability on the structural and functional aspects of macroinvertebrate communities after severe drought

1. Unpredictable, extreme climatic events (e.g. droughts) can potentially destabilize aquatic communities. From 1998 to 2002, southwestern Georgia, U.S.A., experienced the third worst drought of the last 100 years, leading to loss of surface flow in many small streams. We sampled macroinvertebrates, flow and water chemistry in small headwater streams from 2001 to 2007 in two adjacent coastal plain streams of contrasting headwater type (wetland and groundwater‐fed seep) following resumption of flow. 2. Decreasing water temperature, conductivity and nutrient concentrations within the first 2 years of the study indicated flushing of the streambed associated with increased discharge. Invertebrate community composition became less variable over time and during wetter years, reflecting water chemistry, hydrological and climate conditions. 3. A core set of species appeared immediately following breaking of the drought in both streams, reflecting a shared species pool. These species exhibited resilience traits, including short life cycles and resistance to desiccation, which allowed for rapid recovery from disturbance. Such species, which were small‐bodied, sclerotized and abundant in the drift, were then replaced as flows increased by those that were larger, soft‐bodied and rare in drift, suggesting a more stable and less ephemeral habitat. 4. Hydrologic regime and long‐term precipitation indices were strongly correlated with invertebrate community and trait structure. Long‐term data allowed for better interpretation of the effects of infrequent disturbances on aquatic ecosystems. Additionally, long‐term precipitation indices (i.e. 48‐month standardized precipitation index) can indicate the likelihood of a return to drought, allowing for the collection of pre‐disturbance data.     

A 7000‐year history of changing plant trait composition in an Amazonian landscape; the role of humans and climate

Tropical forests are shifting in species and trait composition, but the main underlying causes remain unclear because of the short temporal scales of most studies. Here, we develop a novel approach by linking functional trait data with 7000 years of forest dynamics from a fossil pollen record of Lake Sauce in the Peruvian Amazon. We evaluate how climate and human disturbances affect community trait composition. We found weak relationships between environmental conditions and traits at the taxon level, but strong effects for community‐mean traits. Overall, community‐mean traits were more responsive to human disturbances than to climate change; human‐induced erosion increased the dominance of dense‐wooded, non‐zoochorous species with compound leaves, and human‐induced fire increased the dominance of tall, zoochorous taxa with large seeds and simple leaves. This information can help to enhance our understanding of forest responses to past environmental changes, and improve predictions of future changes in tropical forest composition.     

Long-term seasonal variation in the biological traits of benthic-macroinvertebrates in two Mediterranean-climate streams in California, U.S.A.

1. We investigated the seasonal variation of biological traits and the influence of interannual rainfall variability on this pattern. Using long‐term survey data (6–19 years) from an intermittent and a perennial stream in the Mediterranean‐climate region of northern California, we examined 16 fuzzy‐coded biological traits (e.g. maximum size, life cycle duration, and mode of respiration). 2. Seasonal habitat variability is higher in the intermittent stream than in the perennial stream. During the winter and spring wet‐season both streams flood; however, during the summer dry‐season, the intermittent stream forms isolated pools in (occasionally drying completely). 3. Seasonal habitat variability influenced both taxonomic and biological trait composition. Distinct taxonomic communities were present in each season, particularly in the intermittent stream. The intermittent stream also exhibited more seasonal variation in biological traits than the perennial stream. 4. Despite statistically significant seasonal variation, trait composition was relatively stable among seasons in comparison with taxonomic composition and abundance. Taxonomic composition varied considerably between seasons, because of high seasonal and interannual replacement of taxa resulting from seasonal habitat changes. 5. The seasonality of taxonomic composition and abundance was sensitive to interannual rainfall variability. In dry years, the taxonomic composition of communities was more similar between seasons than in wet years, while trait composition was relatively insensitive to rainfall variability. 6. Despite high seasonal variation in abundance and taxonomic composition, biological traits of aquatic macroinvertebrates varied less and exhibited seasonal stability, which may be a result of the unpredictability and harshness of stream environments.     

Broad‐scale geographic patterns in body size and hind wing development of western Palaearctic carabid beetles(Coleoptera: Carabidae)

Research into large‐scale ecological rules has a long tradition but has received increasing attention over the last two decades. Whereas environmental, especially climatic, influences on the geographic distribution of species traits such as body size are well understood in mammals and birds, our knowledge of the determinants and mechanisms which shape spatial patterns in invertebrate traits is still limited. This study analyzes macroecological patterns in two traits of the highly diverse invertebrate taxon of carabid beetles: body size and hind wing development, the latter being directly linked to species’ dispersal abilities. We tested for potential impacts of environmental variables (spatial, areal, topographic and climate‐related) representing both contemporary conditions and historical processes on large‐scale patterns in the two traits. Regression models revealed hump‐shaped relationships with latitude for both traits in the categories 1) all species, 2) widespread and 3) endemic (restricted‐range) species: body size and the proportion of flightless species increased from northern towards southern Europe and then decreased towards North Africa. The shared and independent influence of environmental factors was analyzed by variation partitioning. While contemporary environmental productivity and stability (represented by measures of ambient energy and water energy balance) had strong positive relationships with carabid body size, patterns in hind wing development were most notably influenced by topography (elevation range). Regions with high elevation range and low historical climate variability (since the last ice age), which likely offer long‐term stable habitats (i.e. glacial refugia), coincide with regions with high proportions of flightless species. Thus geographic patterns in carabid traits tend to be formed not only by recent climate but also by dispersal and historical climate and processes (i.e. glaciations and postglacial colonization).     

Extreme drought pushes stream invertebrate communities over functional thresholds

Functional traits are increasingly being used to predict extinction risks and range shifts under long‐term climate change scenarios, but have rarely been used to study vulnerability to extreme climatic events, such as supraseasonal droughts. In streams, drought intensification can cross thresholds of habitat loss, where marginal changes in environmental conditions trigger disproportionate biotic responses. However, these thresholds have been studied only from a structural perspective, and the existence of functional nonlinearity remains unknown. We explored trends in invertebrate community functional traits along a gradient of drought intensity, simulated over 18 months, using mesocosms analogous to lowland headwater streams. We modelled the responses of 16 traits based on a priori predictions of trait filtering by drought, and also examined the responses of trait profile groups (TPGs) identified via hierarchical cluster analysis. As responses to drought intensification were both linear and nonlinear, generalized additive models (GAMs) were chosen to model response curves, with the slopes of fitted splines used to detect functional thresholds during drought. Drought triggered significant responses in 12 (75%) of the a priori‐selected traits. Behavioural traits describing movement (dispersal, locomotion) and diet were sensitive to moderate‐intensity drought, as channels fragmented into isolated pools. By comparison, morphological and physiological traits showed little response until surface water was lost, at which point we observed sudden shifts in body size, respiration mode and thermal tolerance. Responses varied widely among TPGs, ranging from population collapses of non‐aerial dispersers as channels fragmented to irruptions of small, eurythermic dietary generalists upon extreme dewatering. Our study demonstrates for the first time that relatively small changes in drought intensity can trigger disproportionately large functional shifts in stream communities, suggesting that traits‐based approaches could be particularly useful for diagnosing catastrophic ecological responses to global change.     

Detrimental effects of a novel flow regime on the functional trajectory of an aquatic invertebrate metacommunity

Novel flow regimes resulting from dam operations and overallocation of freshwater resources are an emerging consequence of global change. Yet, anticipating how freshwater biodiversity will respond to surging flow regime alteration requires overcoming two challenges in environmental flow science: shifting from local to riverscape‐level understanding of biodiversity dynamics, and from static to time‐varying characterizations of the flow regime. Here, we used time‐series methods (wavelets and multivariate autoregressive models) to quantify flow‐regime alteration and to link time‐varying flow regimes to the dynamics of multiple local communities potentially connected by dispersal (i.e., a metacommunity). We studied the Chattahoochee River below Buford dam (Georgia, U.S.A.), and asked how flow regime alteration by a large hydropower dam may control the long‐term functional trajectory of the downstream invertebrate metacommunity. We found that seasonal variation in hydropeaking synchronized temporal fluctuations in trait abundance among the flow‐altered sites. Three biological trait states describing adaptation to fast flows benefitted from flow management for hydropower, but did not compensate for declines in 16 “loser” traits. Accordingly, metacommunity‐wide functional diversity responded negatively to hydropeaking intensity, and stochastic simulations showed that the risk of functional diversity collapse within the next 4 years would decrease by 17% if hydropeaking was ameliorated, or by 9% if it was applied every other season. Finally, an analysis of 97 reference and 23 dam‐affected river sites across the U.S. Southeast suggested that flow variation at extraneous, human‐relevant scales (12‐hr, 24‐hr, 1‐week) is relatively common in rivers affected by hydropower dams. This study advances the notion that novel flow regimes are widespread, and simplify the functional structure of riverine communities by filtering out taxa with nonadaptive traits and by spatially synchronizing their dynamics. This is relevant in the light of ongoing and future hydrologic alteration due to climate non‐stationarity and the new wave of dams planned globally.     

Modelling the impact of climate and land use change on the geographical distribution of leaf anatomy in a temperate flora

Variation in plant functional traits has been related to variation in environmental conditions. In particular, the relationship between leaf traits and climate has received much attention. This paper presents a functional‐trait‐centred approach to identify potential impacts of climate and land use change on plant species assemblages. Using species atlas data, we modelled the relative frequencies of species with different leaf anatomies (LARF) as a function of observed climate and land use data on a regular spatial grid across Germany. Subsequently, we projected the geographical distribution of LARF with simulated climate and land use data for the late 21st century under two future scenarios. We used a conditional autoregressive regression model to account for spatially structured variation in LARF that remained unexplained by the environmental factors considered. We found a clear relationship between the climatic gradient of water availability and shifts in LARF: increasing water deficit was associated with a decreasing proportion of species with hygromorphic leaves in the composition and increasing proportions of species with scleromorphic and mesomorphic leaves. The variation in LARF due to land use was only small. Under future environmental scenarios the proportion of species with hygromorphic leaves was projected to decrease in all parts of Germany, whereas the proportions of species with sclero‐ and mesomorphic leaves were projected to increase on average. In particular, Germany's south‐western and north‐eastern areas were projected to experience functional change in LARF. Our study highlights the relationship between functional traits and plant species vulnerability to climate change. Our results suggest that the functional‐trait‐centred approach can provide a powerful additional modelling tool to estimate potential impacts of climate change on plant species assemblages.     

Climatic and trophic processes drive long‐term changes in functional diversity of freshwater invertebrate communities

When investigating the fields of biogeography and macroecology, climate‐ and productivity‐related variables are frequently identified as the strongest correlates of species‐diversity patterns. These variables have been usually merged under the climate/productivity hypothesis and describe the direct and indirect actions of climate on species. Being among the most vulnerable ecosystems to climate change, streams and rivers are expected to be influenced both by climatic and trophic (i.e. productivity‐related) factors. We propose here to distinguish the relative influence of the two processes on large‐scale, long‐term changes in the functional diversity of freshwater invertebrate communities over two decades in France. To this end, we designed two functional indices using invertebrate traits to surrogate the respective mechanisms: climate vulnerability and feeding specialisation. Using geographically weighted regression (GWR) models, we showed that trends in both indices, along with the initial regional species‐pools, have significantly contributed to the overall long‐term increase in functional diversity of invertebrate communities. In addition, we highlighted a strong geographical differentiation in the contribution patterns with the climate vulnerability effect decreasing with latitude and the feeding specialisation effect being higher in headwaters than in large rivers. Finally, taking into account this non‐stationarity in the ecological processes and responses using GWR models allowed explaining about 75% of the long‐term changes in the community diversity. Consequently, this study offers sound perspectives in predicting the future patterns of trends in functional diversity of communities under different scenarios of environmental changes, like climate and/or land‐use.     

Diurnal variation around an optimum and near‐critically high temperature does not alter the performance of an ectothermic aquatic grazer

The growing threat of global climate change has led to a profusion of studies examining the effects of warming on biota. Despite the potential importance of natural variability such as diurnal temperature fluctuations, most experimental studies on warming are conducted under stable temperatures. Here, we investigated whether the responses of an aquatic invertebrate grazer (Lymnaea stagnalis) to an increased average temperature differ when the thermal regime is either constant or fluctuates diurnally. Using thermal response curves for several life‐history and immune defense traits, we first identified the optimum and near‐critically high temperatures that Lymnaea potentially experience during summer heat waves. We then exposed individuals that originated from three different populations to these two temperatures under constant or fluctuating thermal conditions. After 7 days, we assessed growth, reproduction, and two immune parameters (phenoloxidase‐like activity and antibacterial activity of hemolymph) from each individual. Exposure to the near‐critically high temperature led to increased growth rates and decreased antibacterial activity of hemolymph compared to the optimum temperature, whilst temperature fluctuations had no effect on these traits. The results indicate that the temperature level per se, rather than the variability in temperature was the main driver altering trait responses in our study species. Forecasting responses in temperature‐related responses remains challenging, due to system‐specific properties that can include intraspecific variation. However, our study indicates that experiments examining the effects of warming using constant temperatures can give similar predictions as studies with fluctuating thermal dynamics, and may thus be useful indicators of responses in nature.     

A meta‐analysis of the agents of selection on floral traits

Floral traits are hypothesized to evolve primarily in response to selection by pollinators. However, selection can also be mediated by other environmental factors. To understand the relative importance of pollinator‐mediated selection and its variation among trait and pollinator types, we analyzed directional selection gradients on floral traits from experiments that manipulated the environment to identify agents of selection. Pollinator‐mediated selection was stronger than selection by other biotic factors (e.g., herbivores), but similar in strength to selection by abiotic factors (e.g., soil water), providing partial support for the hypothesis that floral traits evolve primarily in response to pollinators. Pollinator‐mediated selection was stronger on pollination efficiency traits than on other trait types, as expected if efficiency traits affect fitness via interactions with pollinators, but other trait types also affect fitness via other environmental factors. In addition to varying among trait types, pollinator‐mediated selection varied among pollinator taxa: selection was stronger when bees, long‐tongued flies, or birds were the primary visitors than when the primary visitors were Lepidoptera or multiple animal taxa. Finally, reducing pollinator access to flowers had a relatively small effect on selection on floral traits, suggesting that anthropogenic declines in pollinator populations would initially have modest effects on floral evolution.     

Afforestation driving long‐term surface water browning

Increase in surface water color (browning), caused by rising dissolved organic carbon (DOC) and iron concentrations, has been widely reported and studied in the last couple of decades. This phenomenon has implications to aquatic ecosystem function and biogeochemical carbon cycling. While recovery from acidification and changes in climate‐related variables, such as precipitation and length of growing season, are recognized as drivers behind browning, land‐use change has received less attention. In this study, we include all of the above factors and aim to discern their individual and combined contribution to water color variation in an unprecedentedly long (1940–2016) and highly resolved dataset (~20 times per month), from a river in southern Sweden. Water color showed high seasonal variability and a marked long‐term increase, particularly in the latter half of the dataset (~1980). Short‐term and seasonal variations were best explained by precipitation, with temperature playing a secondary role. All explanatory variables (precipitation, temperature, S deposition, and land‐use change) contributed significantly and together predicted 75% of the long‐term variation in water color. Long‐term change was best explained by a pronounced increase in Norway spruce (Picea abies Karst) volume—a measure of land‐use change and a proxy for buildup of organic soil layers—and by change in atmospheric S deposition. When modeling water color with a combination of explanatory variables, Norway spruce showed the highest contribution to explaining long‐term variability. This study highlights the importance of considering land‐use change as a factor behind browning and combining multiple factors when making predictions in water color and DOC.     

Environmental,spatial and phylogenetic determinants of fish life‐history traits and functional composition of Australian rivers

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Impacts of multiple anthropogenic stressors on stream macroinvertebrate community composition and functional diversity

Ensuring the provision of essential ecosystem services in systems affected by multiple stressors is a key challenge for theoretical and applied ecology. Trait‐based approaches have increasingly been used in multiple‐stressor research in freshwaters because they potentially provide a powerful method to explore the mechanisms underlying changes in populations and communities. Individual benthic macroinvertebrate traits associated with mobility, life history, morphology, and feeding habits are often used to determine how environmental drivers structure stream communities. However, to date multiple‐stressor research on stream invertebrates has focused more on taxonomic than on functional metrics. We conducted a fully crossed, 4‐factor experiment in 64 stream mesocosms fed by a pristine montane stream (21 days of colonization, 21 days of manipulations) and investigated the effects of nutrient enrichment, flow velocity reduction and sedimentation on invertebrate community, taxon, functional diversity and trait variables after 2 and 3 weeks of stressor exposure. 89% of the community structure metrics, 59% of the common taxa, 50% of functional diversity metrics, and 79% of functional traits responded to at least one stressor each. Deposited fine sediment and flow velocity reduction had the strongest impacts, affecting invertebrate abundances and diversity, and their effects translated into a reduction of functional redundancy. Stressor effects often varied between sampling occasions, further complicating the prediction of multiple‐stressor effects on communities. Overall, our study suggests that future research combining community, trait, and functional diversity assessments can improve our understanding of multiple‐stressor effects and their interactions in running waters.     

The influence of range‐wide plant genetic variation on soil invertebrate communities

Plant genetic variation can have far‐reaching effects on associated communities and ecosystems. Heritable variation in ecologically relevant plant traits is often non‐randomly distributed across a species’ range and can exhibit geographic clines. In the event of range expansions and migration, previously unfamiliar genotypes may have large impacts on resident communities and ecosystems due to the introduction of novel and heritable phenotypic variation. Here we test the hypothesis that geographic origin of a focal plant genotype has effects on belowground invertebrate communities using a common garden field experiment. We sampled soil invertebrates from 103 Oenothera biennis genotypes, which were collected from across the species’ range and planted into a common garden field experiment at the northern range limit. We enumerated 24 000 individuals from 190 morphospecies and found that the diversity, abundance, and composition of soil invertebrate communities varied greatly among plant genotypes. Despite strong effects of plant genotype, we found few genetic correlations between plant traits and soil invertebrate community variables. However, herbivore damage was strongly related to variation in the soil invertebrate community. Geographic origin of plant genotypes had at most a weak effect on belowground communities. We speculate that predicting the extended effects of population movement on associated communities will require detailed knowledge of the trait variation occurring within focal species across particular environmental gradients.     

Limitations of trait‐based approaches for stressor assessment: The case of freshwater invertebrates and climate drivers

The appeal of trait‐based approaches for assessing environmental vulnerabilities arises from the potential insight they provide into the mechanisms underlying the changes in populations and community structure. Traits can provide ecologically based explanations for observed responses to environmental changes, along with predictive power gained by developing relationships between traits and environmental variables. Despite these potential benefits, questions remain regarding the utility and limitations of these approaches, which we explore focusing on the following questions: (a) How reliable are predictions of biotic responses to changing conditions based on single trait–environment relationships? (b) What factors constrain detection of single trait–environment relationships, and how can they be addressed? (c) Can we use information on meta‐community processes to reveal conditions when assumptions underlying trait‐based studies are not met? We address these questions by reviewing published literature on aquatic invertebrate communities from stream ecosystems. Our findings help to define factors that influence the successful application of trait‐based approaches in addressing the complex, multifaceted effects of changing climate conditions on hydrologic and thermal regimes in stream ecosystems. Key conclusions are that observed relationships between traits and environmental stressors are often inconsistent with predefined hypotheses derived from current trait‐based thinking, particularly related to single trait–environment relationships. Factors that can influence findings of trait‐based assessments include intercorrelations of among traits and among environmental variables, spatial scale, strength of biotic interactions, intensity of habitat disturbance, degree of abiotic stress, and methods of trait characterization. Several recommendations are made for practice and further study to address these concerns, including using phylogenetic relatedness to address intercorrelation. With proper consideration of these issues, trait‐based assessment of organismal vulnerability to environmental changes can become a useful tool to conserve threatened populations into the future.