Effects of re‐oligotrophication and climate change on lake thermal structure

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The ecology of the planktonic diatom Cyclotella and its implications for global environmental change studies

The fossil record of diatoms in lake sediments can be used to assess the effects of climate variability on lake ecosystems if ecological relationships between diatom community structure and environmental parameters are well understood. Cyclotella sensu lato taxa are a key group of diatoms that are frequently dominant members of phytoplankton communities in low‐ to moderate‐productivity lakes. Their relative abundances have fluctuated significantly in palaeolimnological records spanning over a century in arctic, alpine, boreal and temperate lakes. This suggests that these species are sensitive to environmental change and may serve as early indicators of ecosystem effects of global change. Yet patterns of change in Cyclotella species are not synchronous or unidirectional across, or even within, regions, raising the question of how to interpret these widespread changes in diatom community structure. We suggest that the path forward in resolving seemingly disparate records is to identify clearly the autecology of Cyclotella species, notably the role of nutrients, dissolved organic carbon and light, coupled with better consideration of both the mechanisms controlling lake thermal stratification processes and the resulting effects of changing lake thermal regimes on light and nutrients. Here we begin by reviewing the literature on the resource requirements of common Cyclotella taxa, illustrating that many studies reveal the importance of light, nitrogen, phosphorus, and interactions among these resources in controlling relative abundances. We then discuss how these resource requirements can be linked to shifts in limnological processes driven by environmental change, including climate‐driven change in lakewater temperature, thermal stratification and nutrient loading, as well as acidification‐driven shifts in nutrients and water clarity. We examine three case studies, each involving two lakes from the same region that have disparate trends in the relative abundances of the same species, and illustrate how the mechanisms by which these species abundances are changing can be deciphered. Ultimately, changes in resource availability and water clarity are key factors leading to shifts in Cyclotella abundances. Tighter integration of the autecology of this important group of diatoms with environmental change and subsequent alterations in limnological processes will improve interpretations of palaeolimnological records, and clarify the drivers of seemingly disparate patterns in fossil records showing widespread and rapid changes across the northern hemisphere.     

Demographic responses of eastern bluebirds to climatic variability in northeastern Arkansas

As climate change continues to alter temperature and precipitation patterns, numerous species have declined. However, populations of some species that show responses to climate change, such as eastern bluebirds (Sialia sialis), have increased or remained stable nationwide. To understand how species are adapting to climate change, we estimated demographic parameters and their responses to climatic variability, using nesting and banding-recapture data between 2003 and 2018 in a northeastern Arkansas eastern bluebird population. Increasing variability in precipitation in the nonbreeding season negatively affected hatchability. Hatching success was negatively affected by increasing variability in maximum temperatures and the number of hot days during the breeding season, but positively affected by increasing winter snow depth. Adult survival was positively affected by increasing snow depth and variability in the number of hot days during the breeding season, but negatively affected by increasing variability in nonbreeding season temperatures. Our results demonstrate that for this study population, annual breeding parameters, though canalized against interannual environmental variation, were affected by seasonal climatic variability. Although climate change may benefit bluebird survival due to increasing variability in winter temperatures and the number of hot days, climatic variability negatively affected breeding parameters and is expected to increase. Because breeding parameters are typically the drivers of population growth rate in short-lived species, these results raise concern for the future of this population of eastern bluebirds.     

Climatic effects on the phenology of lake processes

Populations living in seasonal environments are exposed to systematic changes in physical conditions that restrict the growth and reproduction of many species to only a short time window of the annual cycle. Several studies have shown that climate changes over the latter part of the 20th century affected the phenology and population dynamics of single species. However, the key limitation to forecasting the effects of changing climate on ecosystems lies in understanding how it will affect interactions among species. We investigated the effects of climatic and biotic drivers on physical and biological lake processes, using a historical dataset of 40 years from Lake Washington, USA, and dynamic time‐series models to explain changes in the phenological patterns among physical and biological components of pelagic ecosystems. Long‐term climate warming and variability because of large‐scale climatic patterns like Pacific decadal oscillation (PDO) and El Niño–southern oscillation (ENSO) extended the duration of the stratification period by 25 days over the last 40 years. This change was due mainly to earlier spring stratification (16 days) and less to later stratification termination in fall (9 days). The phytoplankton spring bloom advanced roughly in parallel to stratification onset and in 2002 it occurred about 19 days earlier than it did in 1962, indicating the tight connection of spring phytoplankton growth to turbulent conditions. In contrast, the timing of the clear‐water phase showed high variability and was mainly driven by biotic factors. Among the zooplankton species, the timing of spring peaks in the rotifer Keratella advanced strongly, whereas Leptodiaptomus and Daphnia showed slight or no changes. These changes have generated a growing time lag between the spring phytoplankton peak and zooplankton peak, which can be especially critical for the cladoceran Daphnia. Water temperature, PDO, and food availability affected the timing of the spring peak in zooplankton. Overall, the impact of PDO on the phenological processes were stronger compared with ENSO. Our results highlight that climate affects physical and biological processes differently, which can interrupt energy flow among trophic levels, making ecosystem responses to climate change difficult to forecast.     

Stream diatom assemblages as predictors of climate

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Growth of Blue-Green Algae in the Saidenbach Reservoir and its Relationship to the Silicon Budget

The phytoplankton succession during the summer in the mesotrophic reservoir Saidenbach since 1975 may well be explained by the resource ratio hypothesis. Until 1980, only phosphorus controlled the phytoplankton growth, and diatoms prevailed, because an excesses of silicon existed. From 1981 to 1986, the ratio Si:P often was smaller than 90, a value, critical for the development of the diatom Fragilaria crotonensis. Its reduced growth caused an increased occurrence of blue-greens (mostly Aphanothece clathrata) immediately after the diatom mass development. During these years at first silicon limited phytoplankton growth in summer, later on the growth again was limited by phosphorus. Because of increased Si and P load since 1987 a simultaneous limitation of both nutrients occurs. This leads now to parallel mass developments of diatoms and blue-greens. In order to maintain the positive effect of diatoms (phosphorus transport into the sediment), it is to guarantee a sufficiently high Si:P ratio. If a reduction of P load isn't possible, Si remobilization from the sediment could be increased by artificial changes of the water level.     

Environmental variables drive spatial patterns of trophic diversity in mammals

Understanding environmental drivers of species diversity has become increasingly important under climate change. Different trophic groups (predators, omnivores and herbivores) interact with their environments in fundamentally different ways and may therefore be influenced by different environmental drivers. Using random forest models, we identified drivers of terrestrial mammals' total and proportional species richness within trophic groups at a global scale. Precipitation seasonality was the most important predictor of richness for all trophic groups. Richness peaked at intermediate precipitation seasonality, indicating that moderate levels of environmental heterogeneity promote mammal richness. Gross primary production (GPP) was the most important correlate of the relative contribution of each trophic group to total species richness. The strong relationship with GPP demonstrates that basal-level resource availability influences how diversity is structured among trophic groups. Our findings suggest that environmental characteristics that influence resource temporal variability and abundance are important predictors of terrestrial mammal richness at a global scale.     

Latitudinal patterns of forest ecosystem stability across spatial scales as affected by biodiversity and environmental heterogeneity

Our planet is facing a variety of serious threats from climate change that are unfolding unevenly across the globe. Uncovering the spatial patterns of ecosystem stability is important for predicting the responses of ecological processes and biodiversity patterns to climate change. However, the understanding of the latitudinal pattern of ecosystem stability across scales and of the underlying ecological drivers is still very limited. Accordingly, this study examines the latitudinal patterns of ecosystem stability at the local and regional spatial scale using a natural assembly of forest metacommunities that are distributed over a large temperate forest region, considering a range of potential environmental drivers. We found that the stability of regional communities (regional stability) and asynchronous dynamics among local communities (spatial asynchrony) both decreased with increasing latitude, whereas the stability of local communities (local stability) did not. We tested a series of hypotheses that potentially drive the spatial patterns of ecosystem stability, and found that although the ecological drivers of biodiversity, climatic history, resource conditions, climatic stability, and environmental heterogeneity varied with latitude, latitudinal patterns of ecosystem stability at multiple scales were affected by biodiversity and environmental heterogeneity. In particular, α diversity is positively associated with local stability, while β diversity is positively associated with spatial asynchrony, although both relationships are weak. Our study provides the first evidence that latitudinal patterns of the temporal stability of naturally assembled forest metacommunities across scales are driven by biodiversity and environmental heterogeneity. Our findings suggest that the preservation of plant biodiversity within and between forest communities and the maintenance of heterogeneous landscapes can be crucial to buffer forest ecosystems at higher latitudes from the faster and more intense negative impacts of climate change in the future.     

Long‐term successional forest dynamics: species and community responses to climatic variability

Question: Are trees sensitive to climatic variability, and do tree species differ in their responses to climatic variability? Does sensitivity of forest communities to climatic variability depend on stand composition? Location: Mixed young forest at Walker Branch Watershed near Oak Ridge, East Tennessee, USA. Methods: Using a long‐term dataset (1967–2006), we analyzed temporal forest dynamics at the tree and species level, and community dynamics for forest stands that differed in initial species composition (i.e., chestnut oak, oak–hickory, pine, and yellow poplar stands). Using summer drought and growing season temperature as defined climate drivers, we evaluated relationships between forest dynamics and climate across levels of organization. Results: Over the four‐decade study period, forest communities underwent successional change and substantially increased in biomass. Variation in summer drought and growing season temperature contributed to temporal biomass dynamics for some tree species, but not for others. Stand‐level responses to climatic variability were related to the responses of component species, except in pine stands. Pinus echinata, the dominant species in pine stands, decreased over time due to periodic outbreaks of pine bark beetle (Dendroctonus frontalis). These outbreaks at Walker Branch could not be directly related to climatic conditions. Conclusions: The results indicate that sensitivity of developing forests to climatic variability is stand type‐dependent, and hence is a function of species composition. However, in the long term, direct effects of climatic variability on forest dynamics may be small relative to autogenic successional processes or climate‐related insect outbreaks. Empirical studies testing for interactions between forest succession and climatic variability are needed.     

The effect of climate on the phenology, acorn crop and radial increment of pedunculate oak (Quercus robur) in the middle Volga region, Tatarstan, Russia

Our data, collected in the extreme east of Europe, show that a significant biological effect of climate change has been experienced even in territories where temperature increase has been the lowest. This study documents the climatic response of pedunculate oak (Quercus robur) growing near its north-eastern limits in Europe. It demonstrates the potential of oak trees in old-growth forest to act as proxy climate indicators. Many factors may influence the temporal stability of the growth-climate, acorn crop-climate and first leafing-climate relationships. Climate data, climatic fluctuations, reproduction, genetics and tree-age may relate to this instability. Our results stress that an increase in climate variability or climatic warming resulting from warmer winters or summers could affect the oak population in eastern Europe in a similar way to that in western Europe. These findings, from remnants of oak forest in the middle Volga region of Russia, allow a further understanding of how species could be affected by future climates.     

Ecological and methodological drivers of non-stationarity in tree growth response to climate

Radial tree growth is sensitive to environmental conditions, making observed growth increments an important indicator of climate change effects on forest growth. However, unprecedented climate variability could lead to non-stationarity, that is, a decoupling of tree growth responses from climate over time, potentially inducing biases in climate reconstructions and forest growth projections. Little is known about whether and to what extent environmental conditions, species, and model type and resolution affect the occurrence and magnitude of non-stationarity. To systematically assess potential drivers of non-stationarity, we compiled tree-ring width chronologies of two conifer species, Picea abies and Pinus sylvestris, distributed across cold, dry, and mixed climates. We analyzed 147 sites across the Europe including the distribution margins of these species as well as moderate sites. We calibrated four numerical models (linear vs. non-linear, daily vs. monthly resolution) to simulate growth chronologies based on temperature and soil moisture data. Climate–growth models were tested in independent verification periods to quantify their non-stationarity, which was assessed based on bootstrapped transfer function stability tests. The degree of non-stationarity varied between species, site climatic conditions, and models. Chronologies of P. sylvestris showed stronger non-stationarity compared with Picea abies stands with a high degree of stationarity. Sites with mixed climatic signals were most affected by non-stationarity compared with sites sampled at cold and dry species distribution margins. Moreover, linear models with daily resolution exhibited greater non-stationarity compared with monthly-resolved non-linear models. We conclude that non-stationarity in climate–growth responses is a multifactorial phenomenon driven by the interaction of site climatic conditions, tree species, and methodological features of the modeling approach. Given the existence of multiple drivers and the frequent occurrence of non-stationarity, we recommend that temporal non-stationarity rather than stationarity should be considered as the baseline model of climate–growth response for temperate forests.     

Major changes in trophic dynamics in large, deep sub-alpine Lake Maggiore from 1940s to 2002: a high resolution comparative palaeo-neolimnological study

1. We describe the changes in trophic dynamics in Lake Maggiore from c. 1943 to 2002 using subfossil cladoceran data from a high resolution sediment record, long‐term contemporary data series and historical information. During this period the lake went through a eutrophication phase until 1980 followed by oligotrophication. 2. During the eutrophication period a major increase occurred in the abundance of Chydorus sphaericus, the proportion of planktonic cladocerans and total abundance of cladocerans in the sediment. Since 1980 the abundance declined again and subfossil Eubosmina mucro length and contemporary Daphnia body length increased, most probably as a result of higher abundance of invertebrate predators. 3. Changes in the fish stock composition caused by the introduction of exotic fish during the pre‐eutrophication period and a complete ban on fishing because of Dichloro‐diphenil‐ethanes (DDTs) pollution of the lake (during oligotrophication) could also be detected in the community assemblage and size structure of the sediment zooplankton. 4. We found good correspondence between trophic changes inferred from cladoceran subfossils (community composition, size and predation pressure) and contemporary data, suggesting that sediment samples can be used to infer past development in trophic dynamics, including predation by fish and pelagic invertebrates in lakes with scarce neolimnological data. 5. Furthermore, by combining palaeolimnological cladoceran data rarely obtained from contemporary samples (e.g. benthic and plant‐associated cladocerans, mucro length of bosminids) with contemporary data of organisms poorly represented in the sediment record (e.g. remains of Bythotrephes and fishes) a more complete understanding of changes in trophic dynamics was obtained. 6. The detection in the sediments of meteorological events whose effects on zooplankton had been recorded in the long‐term studies also provided evidence that eutrophication tends to override climate signals. 7. We conclude that a combined palaeo‐neolimnological approach can be a powerful tool for elucidating past changes in the trophic dynamics of lakes and the interaction with climate induced changes, not least when high resolution sediment records are available.     

Predator‐prey migration phenologies remain synchronised in a warming catchment

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Problems with using large‐scale oceanic climate indices to compare climatic sensitivities across populations and species

To understand which populations and species are most sensitive to climate change, studies correlate time series of climate variables with those of traits important for population dynamics, and subsequently compare which aspects of a species’ ecology or life‐history best explain variation in climate sensitivity. Often large‐scale oceanic climate indices (LOCIs) are used as a proxy for local climatic drivers, with many studies reporting geographic gradients in climate sensitivity to LOCIs (e.g. suggesting that species living further from the equator are relatively climate sensitive). However, the relationship between LOCIs and local weather variables also varies geographically, raising the possibility that apparent intra‐ and inter‐specific differences in climate sensitivity to LOCIs could also reflect geographic variation in how well LOCIs function as a proxy for local climatic drivers. This hypothesis is rarely tested due to lack of knowledge about the specific local climatic drivers. Here we show, using reproductive and climate data from 16 long‐term population studies of 7 Australian fairy‐wren species (Malurus genus), that the use of LOCIs can result in 1) strong overestimation of the amount of inter‐specific variation in climate sensitivity and 2) spurious patterns, particularly geographic gradients. Consequently a paradox emerges: LOCIs often explain much of the temporal variation in traits important for population dynamics, but the common usage of LOCIs may prevent meaningful intra‐ and inter‐specific comparisons of climate sensitivities over large spatial scales. Our results thus may offer an alternative interpretation of the widely reported geographic gradients in sensitivity to LOCIs. Future progress will likely require better knowledge about the identity and temporal features of local environmental drivers of population dynamics.     

Using diatom life-forms and ecological guilds to assess organic pollution and trophic level in rivers: a case study of rivers in south-eastern France

The European Union’s Water Framework Directive has set a target of achieving good ecological status for all aquatic environments in Europe by 2015. In order to determine the quality of aquatic environments, biological indicators such as diatoms are often used. However, biotic diatom indices can be difficult and time consuming to use because of complexity of species determination. We investigated whether the biological traits of diatoms in rivers (life-forms, size classes and ecological guilds) could be used to assess organic pollution and trophic level. We worked on a data set comprising 315 diatom species, determined at 328 river stations of south-east France and a variety of parameters. The abundances of some biological traits differed significantly between the different organic pollution and trophic levels, particularly stalked diatoms, and the motile and low-profile guilds.     

Loss of trophic complexity in saline prairie lakes as indicated by stable-isotope based community-metrics

Variations in climate, watershed characteristics and lake-internal processes often result in a large variability of food-web complexity in lake ecosystems. Some of the largest ranges in these environmental parameters can be found in lakes across the northern Great Plains as they are characterized by extreme gradients in respect to lake morphometry and water chemistry, with individual parameters often varying over several orders of magnitude. To evaluate the effects of environmental conditions on trophic complexity in prairie lake food-webs, we analyzed carbon and nitrogen stable isotopes of fishes, zooplankton and littoral macroinvertebrates in 20 lakes across southern Saskatchewan. Our two-year study identified very diverse patterns of trophic complexity, with was predominantly associated with among-lake differences. Small but significant temporal effects were also detected, which were predominantly associated with changes in productivity. The most influential parameters related to changes in trophic complexity among lakes were salinity, complexity of fish assemblage, and indicators of productivity (e.g. nutrients, Chl a). Generally, trophic diversity, number of trophic levels, and trophic redundancy were highest in productive freshwater lakes with diverse fish communities. Surprisingly, mesosaline lakes that were characterized by very low or no predation pressure from fishes were not colonized by invertebrate predators as it is often the case in boreal systems; instead, trophic complexity was further reduced. Together, prairie lake food-webs appear to be highly sensitive to changes in salinity and the loss of piscivorous fishes, making freshwater and mesosaline lakes most vulnerable to the impacts of climate variability. This is particularly important as global circulation models predict future climate warming to have disproportionate negative impacts on hydrologic conditions in this area.     

Cumulative weather effects can impact across the whole life cycle

Predicting how species will be affected by future climatic change requires the underlying environmental drivers to be identified. As vital rates vary over the lifecycle, structured population models derived from statistical environment–demography relationships are often used to inform such predictions. Environmental drivers are typically identified independently for different vital rates and demographic classes. However, these rates often exhibit positive temporal covariance, suggesting that vital rates respond to common environmental drivers. Additionally, models often only incorporate average weather conditions during a single, a priori chosen time window (e.g. monthly means). Mismatches between these windows and the period when the vital rates are sensitive to variation in climate decrease the predictive performance of such approaches. We used a demographic structural equation model (SEM) to demonstrate that a single axis of environmental variation drives the majority of the (co)variation in survival, reproduction, and twinning across six age–sex classes in a Soay sheep population. This axis provides a simple target for the complex task of identifying the drivers of vital rate variation. We used functional linear models (FLMs) to determine the critical windows of three local climatic drivers, allowing the magnitude and direction of the climate effects to differ over time. Previously unidentified lagged climatic effects were detected in this well‐studied population. The FLMs had a better predictive performance than selecting a critical window a priori, but not than a large‐scale climate index. Positive covariance amongst vital rates and temporal variation in the effects of environmental drivers are common, suggesting our SEM–FLM approach is a widely applicable tool for exploring the joint responses of vital rates to environmental change.     

Phenological responses in a sycamore–aphid–parasitoid system and consequences for aphid population dynamics: A 20 year case study

Species interactions have a spatiotemporal component driven by environmental cues, which if altered by climate change can drive shifts in community dynamics. There is insufficient understanding of the precise time windows during which inter‐annual variation in weather drives phenological shifts and the consequences for mismatches between interacting species and resultant population dynamics—particularly for insects. We use a 20 year study on a tri‐trophic system: sycamore Acer pseudoplatanus, two associated aphid species Drepanosiphum platanoidis and Periphyllus testudinaceus and their hymenopteran parasitoids. Using a sliding window approach, we assess climatic drivers of phenology in all three trophic levels. We quantify the magnitude of resultant trophic mismatches between aphids and their plant hosts and parasitoids, and then model the impacts of these mismatches, direct weather effects and density dependence on local‐scale aphid population dynamics. Warmer temperatures in mid‐March to late‐April were associated with advanced sycamore budburst, parasitoid attack and (marginally) D. platanoidis emergence. The precise time window during which spring weather advances phenology varies considerably across each species. Crucially, warmer temperatures in late winter delayed the emergence of both aphid species. Seasonal variation in warming rates thus generates marked shifts in the relative timing of spring events across trophic levels and mismatches in the phenology of interacting species. Despite this, we found no evidence that aphid population growth rates were adversely impacted by the magnitude of mismatch with their host plants or parasitoids, or direct impacts of temperature and precipitation. Strong density dependence effects occurred in both aphid species and probably buffered populations, through density‐dependent compensation, from adverse impacts of the marked inter‐annual climatic variation that occurred during the study period. These findings explain the resilience of aphid populations to climate change and uncover a key mechanism, warmer winter temperatures delaying insect phenology, by which climate change drives asynchronous shifts between interacting species.     

Climate drivers of bark beetle outbreak dynamics in Norway spruce forests

Bark beetles are among the most devastating biotic agents affecting forests globally and several species are expected to be favored by climate change. Given the potential interactions of insect outbreaks with other biotic and abiotic disturbances, and the potentially strong impact of changing disturbance regimes on forest resources, investigating climatic drivers of destructive bark beetle outbreaks is of paramount importance. We analyzed 17 time‐series of the amount of wood damaged by Ips typographus, the most destructive pest of Norway spruce forests, collected across 8 European countries in the last three decades. We aimed to quantify the relative importance of key climate drivers in explaining timber loss dynamics, also testing for possible synergistic effects. Local outbreaks shared the same drivers, including increasing summer rainfall deficit and warm temperatures. Large availability of storm‐felled trees in the previous year was also strongly related to an increase in timber loss, likely by providing an alternative source of breeding material. We did not find any positive synergy among outbreak drivers. On the contrary, the occurrence of large storms reduced the positive effect of warming temperatures and rainfall deficit. The large surplus of breeding material likely boosted I. typographus population size above the density threshold required to colonize and kill healthy trees irrespective of other climate triggers. Importantly, we found strong negative density dependence in I. typographus that may provide a mechanism for population decline after population eruptions. Generality in the effects of complex climatic events across different geographical areas suggests that the large‐scale drivers can be used as early warning indicators of increasing local outbreak probability.     

Interannual variability in carbon dioxide fluxes and flux–climate relationships on grazed and ungrazed northern mixed-grass prairie

The annual carbon (C) budget of grasslands is highly dynamic, dependent on grazing history and on effects of interannual variability (IAV) in climate on carbon dioxide (CO₂) fluxes. Variability in climatic drivers may directly affect fluxes, but also may indirectly affect fluxes by altering the response of the biota to the environment, an effect termed ‘functional change’. We measured net ecosystem exchange of CO₂ (NEE) and its diurnal components, daytime ecosystem CO₂ exchange (P_D) and night‐time respiration (R_E), on grazed and ungrazed mixed‐grass prairie in North Dakota, USA, for five growing seasons. Our primary objective was to determine how climatic anomalies influence variability in CO₂ exchange. We used regression analysis to distinguish direct effects of IAV in climate on fluxes from functional change. Functional change was quantified as the improvement in regression on fitting a model in which slopes of flux–climate relationships vary among years rather than remain invariant. Functional change and direct effects of climatic variation together explained about 20% of variance in weekly means of NEE, P_D, and R_E. Functional change accounted for more than twice the variance in fluxes of direct effects of climatic variability. Grazing did not consistently influence the contribution of functional change to flux variability, but altered which environmental variable best explained year‐to‐year differences in flux–climate slopes, reduced IAV in seasonal means of fluxes, lessened the strength of flux–climate correlations, and increased NEE by reducing R_E relatively more than P_D. Most of these trends are consistent with the interpretation that grazing reduced the influence of plants on ecosystem fluxes. Because relationships between weekly values of fluxes and climatic regulators changed annually, year‐to‐year differences in the C balance of these ecosystems cannot be predicted from knowledge of IAV in climate alone.