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Across the Northern hemisphere, chloride concentrations in lakes are increasing from decades of road salt use, threatening aquatic ecosystems. Zooplankton are sensitive to increasing chloride but few studies have considered how additional stressors, such as soft water or increasing lake temperatures interact with the negative effects of chloride. To determine the effects of increasing chloride and temperature on freshwater zooplankton and phytoplankton communities, we conducted a mesocosm experiment using 30 concentrations of chloride from ambient levels to 1500 mg L−1, ambient or warmed (+2.7°C) temperatures, and plankton from a softwater lake. We compared the results with current Canadian Water Quality Guidelines (CWQG) for chronic exposure to chloride. Although warming elicited little response, increasing chloride concentrations led to reduced zooplankton biomass and richness, even at concentrations below the CWQG, and an increase in cyanobacteria abundance, suggesting that policies regulating road salt application should be reevaluated to better protect aquatic ecosystems. [ABSTRACT FROM AUTHOR]

Interannual variation of 45 annually resolved time series of environmental, limnological, and biotic parameters was quantified (1994-2009) in six lakes within 52000 km2 to test the hypothesis that influx of energy (E; as irradiance, heat, wind) varies synchronously among sites and induces temporal coherence in lakes and their food webs, whereas influx of mass (m; as water, solutes, particles) reduces synchrony because local catchments uniquely modify hydrologic inputs. Overall, 82% of parameters exhibited significant (P < 0.05) synchrony (S) estimated as mean pair-wise correlation of Z-transformed time series. Influx of E as atmospheric heat and irradiance was both more highly synchronous and less temporally variable (months-to-decades) than influx of m as summer precipitation, snow, or river discharge. Similarly, S of limnological parameters varied from 0.08 to 0.85, with variables known to be regulated by E influx (ice melt, gas solubility) up to twofold more coherent than those regulated by m inputs (organic solutes). Pairs of variables linked by simple direct mechanisms exhibited similar S values (air temperature and ice melt, nutrients and algae), whereas the coherence of other parameters (water temperature, mixing) was intermediate to that of multiple regulatory agents. Overall, aggregate measures of plankton density varied more coherently among lakes than did constituent taxa. These findings suggest that environmental variability is transmitted to most levels of aquatic ecosystems, but that the precise effects depend on whether E or in fluxes predominate, the coherence of each forcing mechanism, and the strength of linkages between exogenous forcing and lake response. [ABSTRACT FROM AUTHOR]

G. Evelyn Hutchinson proposed that external control by climate limits the fundamental productivity and the possible diversity of ecological communities. These climatic drivers are currently changing as a result of human activity, which may herald a shift in the influence of climate on global ecosystems. Long-term records reveal a reduction in ice cover on northern lakes over the last several centuries. Hence, we explore whether inter-annual climatic variability, represented by ice cover, influences the productivity and diversity of zooplankton communities in long-term datasets for five lakes in Northern Wisconsin. We used a multilevel modeling approach to test three predictions: (1) density will increase, (2) diversity will increase, and (3) community composition will be altered. We found an inverse relationship between ice-off date and annual zooplankton density. Daphnia density, for example, was inversely related to ice-off date, with 10-fold variability across the gradient of ice-off dates in Northern Wisconsin. In contrast, we did not observe a consistent shift in diversity or community structure. Thus, from ice cover records of northern lakes we found support for Hutchinson’s idea that external climatic forces may regulate aquatic productivity; however, the response was numeric and we did not find evidence that lakes moved closer to maximum diversity on an inter-annual scale. [ABSTRACT FROM AUTHOR]

Understanding how disturbance shapes the dynamics of ecological systems is of fundamental importance in ecology. One emerging approach to revealing and appreciating disturbance effects involves examining disturbance-driven changes in the variability of ecological responses. Variability is rarely employed as a response variable to assess the influence of disturbance, but recent studies indicate that it can be an extremely sensitive metric, capturing differences obscured by averaging and conveying important ecological information about underlying causal processes. In this paper, we present a conceptual model to understand and predict the effects of disturbance on variability. The model estimates qualitative changes in variability by considering disturbance extent, frequency and intensity, as well as ecosystem recovery, and thereby captures not only the immediate effects of disturbance but also those that arise over time due to the biotic response to an event. We evaluate how well the model performs by comparing predictions with empirical results from studies examining a wide variety of disturbances and ecosystems, and discuss factors that may modify or even confound predictions. We include a concise guide to characterizing and detecting changes in variability, highlighting the most common and easily applied methods and conclude by describing several future directions for research. By considering variability as a response to disturbance, we gain another metric of fundamental system behaviour, an improved ability to identify organizing features of ecosystems and a better understanding of the predictability of disturbance-driven change – all critical aspects of assessing ecosystem response to disturbance. [ABSTRACT FROM AUTHOR]

The role that climate plays in determining the temporal dynamics of freshwater zooplankton is poorly understood, despite its potential to synchronize population dynamics on a regional basis. We quantify the spatial synchrony of common zooplankton taxa among lakes in the Dorset, Ontario, area and investigate the climatic mechanisms responsible for these patterns at both annual and seasonal scales. We show that the dynamics of several taxa are characterized by similar patterns of abundance among lakes across the region, suggesting a synchronising effect of climate on zooplankton population abundances. Using multiple regression analyses to compare zooplankton dynamics with climatic and in-lake variables, we find that El Niño – Southern Oscillation (ENSO) events and ice cover variability are among the most important predictors of synchrony at annual and seasonal scales, particularly for Leptodiaptomus and Bosmina. Leptodiaptomus, a dominant herbivore in these lakes, had decreased abundances in years with earlier ice-off dates and shorter periods of ice cover. Secchi depth was also synchronous and appeared to respond to regional meteorological factors (spring temperature and precipitation, ice duration). A warming climate with the ability to modify ice and ENSO dynamics, as well as temperature and precipitation patterns, may also alter the structure and dynamics of zooplankton communities in north-temperate lakes. [ABSTRACT FROM AUTHOR]

Insight into the causes and consequences of changes in aquatic biodiversity requires an improved understanding of the nature of the relationships between species richness and ecosystem function over a much longer temporal perspective than we currently possess. We used high-resolution paleoecological records from two prairie lakes to show that diatom species richness (as fossil frustules) was negatively correlated (r² = 0.09-0.24, p < 0.001) with diatom production (as fossil pigments) during the past 2,000 yr. By comparing analyses from intervals of fresh and saline waters, we demonstrate that these significant richness-production relationships arose during freshwater periods (r² = 0.13-0.45, p < 0.001) and could be eliminated (r² < 0.02, p > 0.1) by abiotic disturbances such as droughts. Procrustes analyses of the concordance of species change within freshwater communities and the change in richness--production relationships through time revealed that shifts in diatom community composition could have a large influence in determining the negative relationship between richness and production. Finally, significant correlations (r² = 0.09-0.24, p < 0.0001) between past diatom species richness and ratios of stable isotopes (primarily δ15N) suggested that C and N biogeochemical cycles are also linked to changes in algal biodiversity. Taken together, these analyses suggest that the ongoing disruption of climate and biogeochemical systems by humans may obscure the relationship between aquatic biodiversity and ecosystem function in the future. [ABSTRACT FROM AUTHOR]

Wildfire activity is increasing globally. The resulting smoke plumes can travel hundreds to thousands of kilometers, reflecting or scattering sunlight and depositing particles within ecosystems. Several key physical, chemical, and biological processes in lakes are controlled by factors affected by smoke. The spatial and temporal scales of lake exposure to smoke are extensive and under‐recognized. We introduce the concept of the lake smoke‐day, or the number of days any given lake is exposed to smoke in any given fire season, and quantify the total lake smoke‐day exposure in North America from 2019 to 2021. Because smoke can be transported at continental to intercontinental scales, even regions that may not typically experience direct burning of landscapes by wildfire are at risk of smoke exposure. We found that 99.3% of North America was covered by smoke, affecting a total of 1,333,687 lakes ≥10 ha. An incredible 98.9% of lakes experienced at least 10 smoke‐days a year, with 89.6% of lakes receiving over 30 lake smoke‐days, and lakes in some regions experiencing up to 4 months of cumulative smoke‐days. Herein we review the mechanisms through which smoke and ash can affect lakes by altering the amount and spectral composition of incoming solar radiation and depositing carbon, nutrients, or toxic compounds that could alter chemical conditions and impact biota. We develop a conceptual framework that synthesizes known and theoretical impacts of smoke on lakes to guide future research. Finally, we identify emerging research priorities that can help us better understand how lakes will be affected by smoke as wildfire activity increases due to climate change and other anthropogenic activities. [ABSTRACT FROM AUTHOR]

Presents information on a study of the temporal coherence of annual abundances among populations of freshwater zooplankton in lakes in Ontario, Canada. Use of the intraclass correlation coefficient; Implications of the correlations; Identification of homogeneous and temporally coherent subsets of lakes for each species.

In lakes, the rates of gross primary production (GPP), ecosystem respiration (R), and net ecosystem production (NEP) are often controlled by resource availability. Herein, we explore how catchment vs. within lake predictors of metabolism compare using data from 16 lakes spanning 39°N to 64°N, a range of inflowing streams, and trophic status. For each lake, we combined stream loads of dissolved organic carbon (DOC), total nitrogen (TN), and total phosphorus (TP) with lake DOC, TN, and TP concentrations and high frequency in situ monitoring of dissolved oxygen. We found that stream load stoichiometry indicated lake stoichiometry for C : N and C : P (r2 = 0.74 and r2 = 0.84, respectively), but not for N : P (r2 = 0.04). As we found a strong positive correlation between TN and TP, we only used TP in our statistical models. For the catchment model, GPP and R were best predicted by DOC load, TP load, and load N : P (R2 = 0.85 and R2 = 0.82, respectively). For the lake model, GPP and R were best predicted by TP concentrations (R2 = 0.86 and R2 = 0.67, respectively). The inclusion of N : P in the catchment model, but not the lake model, suggests that both N and P regulate metabolism and that organisms may be responding more strongly to catchment inputs than lake resources. Our models predicted NEP poorly, though it is unclear why. Overall, our work stresses the importance of characterizing lake catchment loads to predict metabolic rates, a result that may be particularly important in catchments experiencing changing hydrologic regimes related to global environmental change. [ABSTRACT FROM AUTHOR]

Altered ecosystem variability is an important ecological response to disturbance yet understanding of how various attributes of disturbance regimes affect ecosystem variability is limited. To improve the framework for understanding the disturbance regime attributes that affect ecosystem variability, we examine how the introduction of stochasticity to disturbance parameters (frequency, severity and extent) alters simulated recovery when compared to deterministic outcomes from a spatially explicit simulation model. We also examine the agreement between results from empirical studies and deterministic and stochastic configurations of the model. We find that stochasticity in disturbance frequency and spatial extent leads to the greatest increase in the variance of simulated dynamics, although stochastic severity also contributes to departures from the deterministic case. The incorporation of stochasticity in disturbance attributes improves agreement between empirical and simulated responses, with 71% of empirical responses correctly classified by stochastic configurations of the model as compared to 47% using the purely deterministic model. By comparison, only 2% of empirical responses were correctly classified by the deterministic model and misclassified by stochastic configurations of the model. These results indicate that stochasticity in the attributes of a disturbance regime alters the patterns and classification of ecosystem variability, suggesting altered recovery dynamics. Incorporating stochastic disturbance processes into models may thus be critical for anticipating the ecological resilience of ecosystems. [ABSTRACT FROM AUTHOR]

In recent years, unexplained declines in lake total phosphorus (TP) concentrations have been observed at northern latitudes (> 42°N latitude) where most of the world's lakes are found. We compiled data from 389 lakes in Fennoscandia and eastern North America to investigate the effects of climate on lake TP concentrations. Synchrony in year‐to‐year variability is an indicator of climatic influences on lake TP, because other major influences on nutrients (e.g., land use change) are not likely to affect all lakes in the same year. We identified significant synchrony in lake TP both within and among different geographic regions. Using a bootstrapped random forest analysis, we identified winter temperature as the most important factor controlling annual TP, followed by summer precipitation. In Fennoscandia, TP was negatively correlated with the winter East Atlantic Pattern, which is associated with regionally warmer winters. Our results suggest that, in the absence of other overriding factors, lake TP and productivity may decline with continued winter warming in northern lakes. [ABSTRACT FROM AUTHOR]

Cyanobacterial blooms have substantial direct and indirect negative impacts on freshwater ecosystems including releasing toxins, blocking light needed by other organisms, and depleting oxygen. There is growing concern over the potential for climate change to promote cyanobacterial blooms, as the positive effects of increasing lake surface temperature on cyanobacterial growth are well documented in the literature; however, there is increasing evidence that cyanobacterial blooms are also being initiated and persisting in relatively cold‐water temperatures (< 15°C), including ice‐covered conditions. In this work, we provide evidence of freshwater cold‐water cyanobacterial blooms, review abiotic drivers and physiological adaptations leading to these blooms, offer a typology of these lesser‐studied cold‐water cyanobacterial blooms, and discuss their occurrence under changing climate conditions. [ABSTRACT FROM AUTHOR]

The salinization of freshwaters is a global threat to aquatic biodiversity. We quantified variation in chloride (Cl−) tolerance of 19 freshwater zooplankton species in four countries to answer three questions: (1) How much variation in Cl− tolerance is present among populations? (2) What factors predict intraspecific variation in Cl− tolerance? (3) Must we account for intraspecific variation to accurately predict community Cl− tolerance? We conducted field mesocosm experiments at 16 sites and compiled acute LC50s from published laboratory studies. We found high variation in LC50s for Cl− tolerance in multiple species, which, in the experiment, was only explained by zooplankton community composition. Variation in species‐LC50 was high enough that at 45% of lakes, community response was not predictable based on species tolerances measured at other sites. This suggests that water quality guidelines should be based on multiple populations and communities to account for large intraspecific variation in Cl− tolerance. [ABSTRACT FROM AUTHOR]

Human-induced salinization caused by the use of road deicing salts, agricultural practices, mining operations, and climate change is a major threat to the biodiversity and functioning of freshwater ecosystems. Yet, it is unclear if freshwater ecosystems are protected from salinization by current water quality guidelines. Leveraging an experimental network of land-based and in-lake mesocosms across North America and Europe, we tested how salinization— indicated as elevated chloride (Cl2) concentration—will affect lake food webs and if two of the lowest Cl2 thresholds found globally are sufficient to protect these food webs. Our results indicated that salinization will cause substantial zooplankton mortality at the lowest Cl2 thresholds established in Canada (120 mg Cl2/L) and the United States (230 mg Cl2/L) and throughout Europe where Cl2 thresholds are generally higher. For instance, at 73% of our study sites, Cl2 concentrations that caused a ≥50% reduction in cladoceran abundance were at or below Cl2 thresholds in Canada, in the United States, and throughout Europe. Similar trends occurred for copepod and rotifer zooplankton. The loss of zooplankton triggered a cascading effect causing an increase in phytoplankton biomass at 47% of study sites. Such changes in lake food webs could alter nutrient cycling and water clarity and trigger declines in fish production. Current Cl2 thresholds across North America and Europe clearly do not adequately protect lake food webs. Water quality guidelines should be developed where they do not exist, and there is an urgent need to reassess existing guidelines to protect lake ecosystems from human-induced salinization. [ABSTRACT FROM AUTHOR]

Carbon and nitrogen stable isotope ratios are increasingly used to study long-term food web change. Temporal variation at the base of the food web may impact the accuracy of trophic niche estimates, but data describing interannual baseline variation are limited. We quantified baseline variation over a 23-year period in a north-temperate lake and used a simulation model to examine how this variation might affect consumer trophic niche estimates. Interannual variation in C and N stable isotope ratios was significant for both benthic and pelagic primary consumer baselines. Long-term linear trends and shorter-term autoregressive patterns were apparent in the data. There were no correlations among benthic and pelagic C and N baselines. Simulations demonstrated that error in estimated fish trophic niches, but not bias, increased substantially when sampling of baselines was incomplete. Accurate trophic niche estimates depended more on accurate estimation of baseline time series than on accurate estimation of growth and turnover rates. These results highlight the importance of previous and continued efforts to constrain bias and error in long-term stable isotope food web studies. [ABSTRACT FROM AUTHOR]

Population dynamics are influenced by drivers acting from outside and from within an ecosystem. Extrinsic forces operating over broad spatial scales can impart synchronous behavior to separate populations, while internal, system-specific drivers often lead to idiosyncratic behavior. Here, we demonstrate synchrony in community-level dynamics among phytoplankton and bacteria in six north temperate humic lakes. The influence of regional meteorological factors explained much of the temporal variability in the phytoplankton community, and resulted in synchronous patterns of community change among lakes. Bacterial dynamics, in contrast, were driven by system-specific interactions with phytoplankton. Despite the importance of intrinsic factors for determining bacterial community composition and dynamics, we demonstrated that biological interactions transmitted the signal of the regional extrinsic drivers to the bacterial communities, ultimately resulting in synchronous community phenologies for bacterioplankton communities as well. This demonstrates how linkages between the components of a complex biological system can work to simplify the dynamics of the system and implies that it may be possible to predict the behavior of microbial communities responsible for important biogeochemical services in the landscape.The ISME Journal (2007) 1, 38–47; doi:10.1038/ismej.2007.6 [ABSTRACT FROM AUTHOR]

The intensity and frequency of storms are projected to increase in many regions of the world because of climate change. Storms can alter environmental conditions in many ecosystems. In lakes and reservoirs, storms can reduce epilimnetic temperatures from wind‐induced mixing with colder hypolimnetic waters, direct precipitation to the lake's surface, and watershed runoff. We analyzed 18 long‐term and high‐frequency lake datasets from 11 countries to assess the magnitude of wind‐ vs. rainstorm‐induced changes in epilimnetic temperature. We found small day‐to‐day epilimnetic temperature decreases in response to strong wind and heavy rain during stratified conditions. Day‐to‐day epilimnetic temperature decreased, on average, by 0.28°C during the strongest windstorms (storm mean daily wind speed among lakes: 6.7 ± 2.7 m s−1, 1 SD) and by 0.15°C after the heaviest rainstorms (storm mean daily rainfall: 21.3 ± 9.0 mm). The largest decreases in epilimnetic temperature were observed ≥2 d after sustained strong wind or heavy rain (top 5th percentile of wind and rain events for each lake) in shallow and medium‐depth lakes. The smallest decreases occurred in deep lakes. Epilimnetic temperature change from windstorms, but not rainstorms, was negatively correlated with maximum lake depth. However, even the largest storm‐induced mean epilimnetic temperature decreases were typically <2°C. Day‐to‐day temperature change, in the absence of storms, often exceeded storm‐induced temperature changes. Because storm‐induced temperature changes to lake surface waters were minimal, changes in other limnological variables (e.g., nutrient concentrations or light) from storms may have larger impacts on biological communities than temperature changes. [ABSTRACT FROM AUTHOR]

We evaluated the crustacean zooplankton size spectrum as an indicator of lake characteristics and ecosystem change. First, we used time series from seven Canadian Shield lakes to identify the factors associated with among-lake and among-year variability in the spectrum slope (relative abundance of small and large zooplankton) and centered height (total abundance). Second, we used time series from an invaded and three control lakes to assess change in mean and variability in slope and height due to a Bythotrephes invasion. We found that the slope and height reflected among-lake predictors related to morphometry. The slope was responsive to long-term declining lake phosphorus levels, whereas the height reflected both increases in dissolved organic carbon and decreases in ice duration. We detected a significant increase (i.e., flattening) in mean slope and substantial (up to 120%) increases in the CV of height after Bythotrephes invaded Harp Lake. Thus, the zooplankton size spectrum was responsive to long-term environmental change, and a strong top-down perturbation can be detected through regular and frequent monitoring programs. [ABSTRACT FROM AUTHOR]

Lakes are traditionally classified based on their thermal regime and trophic status. While this classification adequately captures many lakes, it is not sufficient to understand seasonally ice‐covered lakes, the most common lake type on Earth. We describe the inverse thermal stratification in 19 highly varying lakes and derive a model that predicts the temperature profile as a function of wind stress, area, and depth. The results suggest an additional subdivision of seasonally ice‐covered lakes to differentiate underice stratification. When ice forms in smaller and deeper lakes, inverse stratification will form with a thin buoyant layer of cold water (near 0°C) below the ice, which remains above a deeper 4°C layer. In contrast, the entire water column can cool to ∼0°C in larger and shallower lakes. We suggest these alternative conditions for dimictic lakes be termed "cryostratified" and "cryomictic." Plain Language Summary: Most mid and high latitude lakes are seasonally ice‐covered and have only been classified based on the thermal structure and trophic status during the open‐water season in summer. However, limited temperatures observations in these ice‐covered lakes suggest that there is a wide range of thermal structures over winter. We developed an analytical model to predict the average water temperature at the time of ice formation based on the strength of the surface winds, the area of the lake, and the maximum depth of the lake. Using both the analytical model and water temperature data from 19 different lakes in North America, Europe, and Asia, we found that the time of ice formation in lakes that are large or experience strong winds were later compared to lakes that are small or experience weak winds. The larger and windier lakes are also colder (0°C∼2°C) than smaller and calmer lakes (2°C∼4°C) at the time of ice formation. This suggests that these seasonally ice‐covered lakes can be subdivided into two additional classes during winter. The analytical model and the new categorization have important consequences for understanding fish habitat under the ice and the potential effects of climate change on these seasonally ice‐covered lakes. Key Points: Standard classifications of dimictic lakes do not consider how variable the initial thermal stratification can be under winter lake iceLakes that are shallow or windy can cool to near 0°C–1°C before ice forms and are weakly stratified, which we term "cryomictic"Deeper lakes or those with calmer winds, result in ice forming just above deeper waters of 3°C–4°C, which we term "cryostratified" [ABSTRACT FROM AUTHOR]

Scientific Significance Statement: Metabolic stoichiometry predicts that dissolved oxygen (O2) and carbon dioxide (CO2) in aquatic ecosystems should covary inversely; however, field observations often diverge from theoretical expectations. Here, we propose a suite of metrics describing this O2 and CO2 decoupling and introduce a conceptual framework for interpreting these metrics within aquatic ecosystems. Within this framework, we interpret cross‐system patterns of high‐frequency O2 and CO2 measurements in 11 northern lakes and extract emergent insights into the metabolic behavior and the simultaneous roles of chemical and physical forcing in shaping ecosystem processes. This approach leverages the power of high‐frequency paired O2–CO2 measurements, and yields a novel, integrative aquatic system typology which can also be applicable more broadly to streams and rivers, wetlands and marine systems. [ABSTRACT FROM AUTHOR]

Lentic ecosystems are important agents of local and global carbon cycling, but their contribution varies along gradients of dissolved organic matter (DOM) and productivity. We investigated how contrasting summer and autumn precipitation can shape annual and inter-annual variation in ecosystem carbon (C) flux (gross primary production (GPP), ecosystem respiration (ER), and CO 2 efflux) in two subtropical lakes differing substantially in trophic state and water color. Instrumented buoys recorded time series of free-water DO, terrestrial DOM (tDOM), chlorophyll a , water temperature profiles, and meteorological measurements over five years (2009–2011 and 2014–2015). Reduced precipitation caused immediate and prolonged effects on C flux in both lakes. During the drought year (2014) GPP and ER declined by 60 to 80% and both lakes were either CO 2 sinks or neutral. In the subsequent wet year (2015), GPP and ER increased by 40 to 110%, and both lakes shifted to strong net CO 2 emitters. Higher ecosystem R resulted from larger GPP while higher tDOM contributed to a dramatic increase in dissolved inorganic carbon, which intensified CO 2 emission in both lakes. C flux was more responsive in the clear mesotrophic lake, declining by approximately 40% in the cumulative GPP and ER, and increasing by >400% in CO 2 efflux whereas changes in the oligotrophic colored lake were more modest (approximately 30% and 300% for metabolic declines and efflux increases, respectively). Temporal variation and magnitude of C flux were governed by tDOM-mediated changes in epilimnetic nutrient levels and hypolimnetic light availability. This study demonstrated terrestrial loads of DOM strongly influence the inter-annual response and sensitivity of ecosystem C flux to variation in inter-annual precipitation. Our findings have important implications for predicting the trend, magnitude, duration, and sensitivity of the response of C flux in subtropical lakes/reservoirs to future changes in precipitation patterns under altered climatic conditions. Unlabelled Image • Effects of droughts on inter-annual C fluxes were studied in subtropical lakes. • Multi-parameter probes were deployed to record key parameters over five years. • Lakes shifted from a C sink in dry year to C source in subsequent normal year. • C fluxes were more responsive to droughts in the mesotrophic colored lake. • Terrestrial DOM serves as a driver for the response of lakes to droughts. [ABSTRACT FROM AUTHOR]

Developing the crustacean zooplankton community size spectrum into an indicator of change in lakes requires quantification of the natural variability in the size spectrum related to broad-scale seasonal, annual, and spatial factors. Characterizing seasonal patterns of variation in the size spectrum is necessary so that monitoring programs can be designed to minimize the masking effects that seasonal processes can have on detecting longer-term temporal change. We used a random effects model to measure monthly, annual, and interlake variability in the slope (i.e., relative abundance of small and large organisms) and centered height (i.e., total abundance) of the crustacean zooplankton normalized abundance size spectrum from 1981 to 2011 among eight Canadian Shield lakes. Consistent with theoretical predictions, the slope was a relatively stable characteristic of the zooplankton community compared with the height, which varied significantly among lakes. We identified a seasonal signal in height and slope and used a mixed effects model to characterize the linear rate of change from May to October; there was an overall decline in height and an overall increase in slope. Seasonal variance was greater than annual variance for both the height and the slope, suggesting that long-term monitoring of lakes and interlake comparisons using zooplankton size spectra should be based on temporally standardized sampling protocols that minimize the effects of seasonal processes. We recommend sampling the zooplankton community in midsummer because this results in size spectrum estimates close to seasonal mean values. [ABSTRACT FROM AUTHOR]

Researchers have long viewed patterns of species association as key to understanding the processes that structure communities. Community‐level tests of species association have received the most attention; however, pairwise species associations may offer greater opportunity for linking patterns to specific mechanisms. Although several tests of pairwise association have been developed, there remain gaps in our understanding of their performance. Consequently, it is unclear whether these methods reliably detect patterns of association, or if any one method is superior. We maximized association patterns for single species pairs in synthetic community matrices and examined how accurately five pairwise association tests found that pair, while not finding others (i.e., type I and II error rates). All tests are more likely to miss patterns of association than to falsely detect them. When we maximized association for a species pair that included one or more rare or common species, tests were frequently unable to identify that pair as significantly associated. Consequently, these tests are best suited for identifying significant associations between pairs of species that occur in an intermediate number of samples; for such pairs, three of the five tests considered here detected 100% of the pairs for which we maximized associations. [ABSTRACT FROM AUTHOR]

Heat fluxes at the lake surface play an integral part in determining the energy budget and thermal structure in lakes, including regulating how lakes respond to climate change. We explore patterns in turbulent heat fluxes, which vary across temporal and spatial scales, using in situ high‐frequency monitoring data from 45 globally distributed lakes. Our analysis demonstrates that some of the lakes studied follow a marked seasonal cycle in their turbulent surface fluxes and that turbulent heat loss is highest in larger lakes and those situated at low latitude. The Bowen ratio, which is the ratio of mean sensible to mean latent heat fluxes, is smaller at low latitudes and, in turn, the relative contribution of evaporative to total turbulent heat loss increases toward the tropics. Latent heat transfer ranged from ~ 60% to > 90% of total turbulent heat loss in the examined lakes. The Bowen ratio ranged from 0.04 to 0.69 and correlated significantly with latitude. The relative contributions to total turbulent heat loss therefore differ among lakes, and these contributions are influenced greatly by lake location. Our findings have implications for understanding the role of lakes in the climate system, effects on the lake water balance, and temperature‐dependent processes in lakes. [ABSTRACT FROM AUTHOR]

Abstract: The vertical distribution of chlorophyll in stratified lakes and reservoirs frequently exhibits a maximum peak deep in the water column, referred to as the deep chlorophyll maximum (DCM). DCMs are ecologically important hot spots of primary production and nutrient cycling, and their location can determine vertical habitat gradients for primary consumers. Consequently, the drivers of DCM structure regulate many characteristics of aquatic food webs and biogeochemistry. Previous studies have identified light and thermal stratification as important drivers of summer DCM depth, but their relative importance across a broad range of lakes is not well resolved. We analyzed profiles of chlorophyll fluorescence, temperature, and light during summer stratification from 100 lakes in the Global Lake Ecological Observatory Network (GLEON) and quantified two characteristics of DCM structure: depth and thickness. While DCMs do form in oligotrophic lakes, we found that they can also form in eutrophic to dystrophic lakes. Using a random forest algorithm, we assessed the relative importance of variables associated with light attenuation vs. thermal stratification for predicting DCM structure in lakes that spanned broad gradients of morphometry and transparency. Our analyses revealed that light attenuation was a more important predictor of DCM depth than thermal stratification and that DCMs deepen with increasing lake clarity. DCM thickness was best predicted by lake size with larger lakes having thicker DCMs. Additionally, our analysis demonstrates that the relative importance of light and thermal stratification on DCM structure is not uniform across a diversity of lake types. [ABSTRACT FROM AUTHOR]

Turbulent fluxes across the air-water interface are integral to determining lake heat budgets, evaporation, and carbon emissions from lakes. The stability of the atmospheric boundary layer (ABL) influences the exchange of turbulent energy. We explore the differences in over-lake ABL stability using data from 39 globally distributed lakes. The frequency of unstable ABL conditions varied between lakes from 71 to 100% of the time, with average air temperatures typically several degrees below the average lake surface temperature. This difference increased with decreasing latitude, resulting in a more frequently unstable ABL and a more efficient energy transfer to and from the atmosphere, toward the tropics. In addition, during summer the frequency of unstable ABL conditions decreased with increasing lake surface area. The dependency of ABL stability on latitude and lake size has implications for heat loss and carbon fluxes from lakes, the hydrologic cycle, and climate change effects. [ABSTRACT FROM AUTHOR]

Lakewater calcium (Ca) decline affects softwater lakes across the Canadian Shield. Ca decline is one consequence of acid deposition, and has impeded biological recovery in formerly acidified lakes. Reduced Ca availability may advantage taxa better adapted to low Ca waters. Crosson Lake in south-central Ontario (Canada) has an extensive monitoring record and has experienced strong Ca decline since the late 1970s, recently falling below 1.5 mg L, a threshold value for some large Daphnia O.F. Müller, 1785 taxa. Paleolimnological analysis of the sedimentary cladoceran assemblages revealed that changes associated with the Ca decline began in the early 1970s. Relative abundances of the jelly-clad Holopedium glacialis Rowe, 2007 and the Daphnia pulex complex increased while Ca-sensitive members of the Daphnia longispina complex decreased. Zooplankton net hauls (1981-2010) corroborate the paleolimnological analysis, revealing that increases in the D. pulex complex were due to Daphnia catawba Coker, 1926 (a taxon tolerant of low Ca). Lakewater Ca, dissolved organic carbon and total phosphorus explain a significant amount of variation within the cladoceran community; however, the relationship between Ca concentration and the daphniid community was most apparent. The Crosson Lake paleolimnological and direct monitoring data may describe ecological changes that are also occurring in many other softwater lakes across the Canadian Shield and elsewhere. [ABSTRACT FROM AUTHOR]

Ecological and biogeochemical processes in lakes are strongly dependent upon water temperature. Long-term surface warming of many lakes is unequivocal, but little is known about the comparative magnitude of temperature variation at diel timescales, due to a lack of appropriately resolved data. Here we quantify the pattern and magnitude of diel temperature variability of surface waters using high-frequency data from 100 lakes. We show that the near-surface diel temperature range can be substantial in summer relative to long-term change and, for lakes smaller than 3 km², increases sharply and predictably with decreasing lake area. Most small lakes included in this study experience average summer diel ranges in their near-surface temperatures of between 4 and 7°C. Large diel temperature fluctuations in the majority of lakes undoubtedly influence their structure, function and role in biogeochemical cycles, but the full implications remain largely unexplored. [ABSTRACT FROM AUTHOR]

Multiple natural and anthropogenic stressors pose a serious threat to the health and diversity of biological communities. A growing body of research suggests that one consequence of these stressors has been a regional-scale loss of crayfish populations across south-central Ontario. The mechanisms contributing to the imperilment of crayfish probably include historical acidification and metal contamination, shoreline development and habitat loss, increased water temperature, declining [Ca] in lake water, and invasion by nonnative species (i.e., Smallmouth Bass and the rusty crayfish). Long-term monitoring data of crayfish populations in Algonquin Provincial Park (Ontario) have documented marked declines in Cambarus bartonii abundance despite the relatively isolated nature of the study lakes. Furthermore, monitoring data indicate that the onset of this decline may predate the monitoring record, thereby creating a need for the use of paleolimnological techniques to infer past conditions. We used a multiple proxy approach (i.e., diatoms and cladocerans) to reconstruct key environmental variables (e.g., pH) and long-term changes in lake chemistry (e.g., [Ca]) as they pertain to crayfish population loss. We found that interactions between pH and [Ca] have led to the decline in crayfish abundance and to the continued lack of recovery of these populations. Furthermore, lake-water [Ca] in all 4 Algonquin Park lakes are now <2.0 mg/L, the lowest hypothesized requirement for freshwater crayfish (2–10 mg/L), suggesting that short-term recovery of crayfish populations is unlikely, despite pH recovery. [ABSTRACT FROM AUTHOR]

We quantify the allochthonous organic carbon (OC) budgets for seven north temperate lakes, using diverse information about their land cover, hydrology, and limnological characteristics. We develop a simple equilibrium model within a Bayesian framework that exploits the differences among the lakes to estimate three key rates: aerial loading (AOC) and wetland loading (WOC) from adjacent ecosystems and whole-lake mineralization of OC (RD OC). Combined with observational data, these rates allow for estimates of the total OC loads, mineralization, and sedimentation within lakes and export to downstream ecosystems. AOC was 1.15 g C m-1 (shoreline) d-1, W ranged from 0.72 to 3.00 g C m-1 (shoreline) d-1, and RDOC, normalized to 20°C, ranged from 0.00083 to 0.0015 d-1. Total loads ranged from about 5 to 55 g C m-2 yr-1. Ecosystems immediately adjacent to lakes accounted for one-half or more of total OC loads for some lakes. Whether a lake processed and stored more allochthonous OC than it exported depended primarily on hydrologic residence time. Our equilibrium model provides a parsimonious approach to quantifying allochthonous OC budgets in lakes with relatively minimal baseline data. [ABSTRACT FROM AUTHOR]

The development of complete regional carbon (C) budgets for different biomes is an integral step in the effort to predict global response and potential feedbacks to a changing climate regime. Wetland and lake contributions to regional C cycling remain relatively uncertain despite recent research highlighting their importance. Using a combination of field surveys and tower-based carbon dioxide (CO) flux measurements, modeling, and published literature, we constructed a complete C budget for the Northern Highlands Lake District in northern Wisconsin/Michigan, a ∼6400 km region rich in lakes and wetlands. This is one of the first regional C budgets to incorporate aquatic and terrestrial C cycling under the same framework. We divided the landscape into three major compartments (forests, wetlands, and surface waters) and quantified all major C fluxes into and out of those compartments, with a particular focus on atmospheric exchange but also including sedimentation in lakes and hydrologic fluxes. Landscape C storage was dominated by peat-containing wetlands and lake sediments, which make up only 20% and 13% of the landscape area, respectively, but contain >80% of the total fixed C pool (ca. 400 Tg). We estimated a current regional C accumulation of 1.1±0.1 Tg yr, and the largest regional flux was forest net ecosystem exchange (NEE) into aggrading forests for a total of 1.0±0.1 Tg yr. Mean wetland NEE (0.12±0.06 Tg yr into wetlands), lake CO emissions and riverine efflux (each ca. 0.03±0.01 Tg yr) were smaller but of consequence to the overall budget. Hydrologic transport from uplands/wetlands to surface waters within the region was an important vector of terrestrial C. Regional C fluxes and pools would be misrepresented without inclusion of surface waters and wetlands, and C budgets in heterogeneous landscapes open opportunities to examine the sensitivities of important fluxes to changes in climate and land use/land cover. [ABSTRACT FROM AUTHOR]

Ecology Letters (2010) 13: 453–463 Environmental variability in space and time is a primary mechanism allowing species that share resources to coexist. Fluctuating conditions are a double edged sword for diversity, either promoting coexistence through temporal niche partitioning or excluding species by stochastic extinctions. The net effect of environmental variation on diversity is largely unknown. We examined the association between zooplankton species richness in lakes and environmental variability on interannual, seasonal and shorter time scales, as well as long-term average conditions. We analyzed data on physical, chemical and biological limnology in 53 temperate zone lakes in North America and Europe sampled over a combined 1042 years. Large fluctuations in pH, phosphorus and dissolved organic carbon concentration on different time scales were associated with reduced zooplankton species richness. More species were found in lakes that showed greater temperature variation on all time scales. Environmental variability on different time scales showed similar or, in some cases, stronger associations with zooplankton species richness compared with long-term average conditions. Our results suggest that temporal fluctuations in the chemical environment tend to exclude zooplankton species while temperature variability promotes greater richness. The results indicate that anthropogenic increases in temporal variability of future climates may have profound effects on biodiversity. [ABSTRACT FROM AUTHOR]

Ecology Letters (2010) 13: 162–174 Predicting community and species responses to disturbance is complicated by incomplete knowledge about species traits. A phylogenetic framework should partially solve this problem, as trait similarity is generally correlated with species relatedness, closely related species should have similar sensitivities to disturbance. Disturbance should thus result in community assemblages of closely related species. We tested this hypothesis with 18 disturbed and 16 reference whole-lake, long-term zooplankton data sets. Regardless of disturbance type, communities generally contained more closely related species when disturbed. This effect was independent of species richness, evenness, and abundance. Communities already under stress (i.e., those in acidic lakes) changed most when disturbed. Species sensitivities to specific disturbances were phylogenetically conserved, were independent of body size, and could be predicted by the sensitivities of close relatives within the same community. Phylogenetic relatedness can effectively act as a proxy for missing trait information when predicting community and species responses to disturbance. [ABSTRACT FROM AUTHOR]

Time trends in abundance, body size, species richness, and species composition indicate that crustacean zooplankton communities of southern Canadian Shield lakes changed between 1980 and 2003. Total abundance did not decline despite reductions in total phosphorus, but all other metrics changed. Species richness declined in Harp Lake (Ontario, Canada) following its Bythotrephes invasion, but richness increased in three other lakes. Average cladoceran body length increased from 0.6 to 1.0 mm in seven of the lakes, as larger-bodied taxa replaced smaller ones. Correlations with water quality and fish metrics suggest that cladoceran size increases were attributable to many factors: a decline in food availability following declining phosphorus levels increasing the competitive advantage of larger herbivores, a decline in acidity favouring the larger, acid-sensitive daphniids, and reduced risk of planktivory linked to a rise in dissolved organic carbon levels and changes in predation regimes. Zooplankton communities on the Canadian Shield are changing, and these changes are best viewed in a multiple-stressor context. Key anthropogenic stressors have also changed and may do so again if Ca concentrations continue to decline. [ABSTRACT FROM AUTHOR]

We compared long-term change in two lake districts, one in a forested rural setting and the other in an urbanizing agricultural region, using lakes as sentinel ecosystems. Human population growth and land-use change are important drivers of ecosystem change in both regions. Biotic changes such as habitat loss, species invasions, and poorer fishing were prevalent in the rural region, and lake hydrology and biogeochemistry responded to climate trends and landscape position. Similar biotic changes occurred in the urbanizing agricultural region, where human-caused changes in hydrology and biogeochemistry had conspicuous effects. Feedbacks among ecosystem dynamics, human uses, economics, social dynamics, and policy and practice are fundamental to understanding change in these lake districts. Sustained support for interdisciplinary collaboration is essential to build understanding of regional change. [ABSTRACT FROM AUTHOR]

The provisioning of sustaining goods and services that we obtain from natural ecosystems is a strong economic justification for the conservation of biological diversity. Understanding the relationship between these goods and services and changes in the size, arrangement, and quality of natural habitats is a fundamental challenge of natural resource management. In this paper, we describe a new approach to assessing the implications of habitat loss for loss of ecosystem services by examining how the provision of different ecosystem services is dominated by species from different trophic levels. We then develop a mathematical model that illustrates how declines in habitat quality and quantity lead to sequential losses of trophic diversity. The model suggests that declines in the provisioning of services will initially be slow but will then accelerate as species from higher trophic levels are lost at faster rates. Comparison of these patterns with empirical examples of ecosystem collapse (and assembly) suggest similar patterns occur in natural systems impacted by anthropogenic change. In general, ecosystem goods and services provided by species in the upper trophic levels will be lost before those provided by species lower in the food chain. The decrease in terrestrial food chain length predicted by the model parallels that observed in the oceans following overexploitation. The large area requirements of higher trophic levels make them as susceptible to extinction as they are in marine systems where they are systematically exploited. Whereas the traditional species area curve suggests that 50% of species are driven extinct by an order-of-magnitude decline in habitat abundance, this magnitude of loss may represent the loss of an entire trophic level and all the ecosystem services performed by the species on this trophic level. [ABSTRACT FROM AUTHOR]

The species–time relationship (STR) describes how the species richness of a community increases with the time span over which the community is observed. This pattern has numerous implications for both theory and conservation in much the same way as the species–area relationship (SAR). However, the STR has received much less attention and to date only a handful of papers have been published on the pattern. Here we gather together 984 community time-series, representing 15 study areas and nine taxonomic groups, and evaluate their STRs in order to assess the generality of the STR, its consistency across ecosystems and taxonomic groups, its functional form, and its relationship to local species richness. In general, STRs were surprisingly similar across major taxonomic groups and ecosystem types. STRs tended to be well fit by both power and logarithmic functions, and power function exponents typically ranged between 0.2 and 0.4. Communities with high richness tended to have lower STR exponents, suggesting that factors increasing richness may simultaneously decrease turnover in ecological systems. Our results suggest that the STR is as fundamental an ecological pattern as the SAR, and raise questions about the general processes underlying this pattern. They also highlight the dynamic nature of most species assemblages, and the need to incorporate time scale in both basic and applied research on species richness patterns. [ABSTRACT FROM AUTHOR]

The species-area relationship (SAR) plays a central role in biodiversity research, and recent work has increased awareness of its temporal analogue, the species-time relationship (STR). Here we provide evidence for a general' species-time-area relationship (STAR), in which species number is a function of the area and time span of sampling, as well as their interaction. For eight assemblages, ranging from lake zooplankton to desert rodents, this model outperformed a sampling-based model and two simpler models in which area and time had independent effects. In every case, the interaction term was negative, meaning that rates of species accumulation in space decreased with the time span of sampling, while species accumulation rates in time decreased with area sampled. Although questions remain about its precise functional form, the STAR provides a tool for scaling species richness across time and space, for comparing the relative rates of species turnover in space and time at different scales of sampling, and for rigorous testing of mechanisms proposed to drive community dynamics. Our results show that the SAR and STR are not separate relationships but two dimensions of one unified pattern. [ABSTRACT FROM AUTHOR]

Six high-resolution climatic reconstructions, based on diatom analyses from lake sediment cores from the northern prairies of North America, show that shifts in drought conditions on decadal through multicentennial scales have prevailed in this region for at least the last two millennia. The predominant broad-scale pattern seen at all sites is a major shift in moisture regimes from wet to dry, or vice versa (depending on location), that occurred after a period of relative stability. These large-scale shifts at the different sites exhibit spatial coherence at regional scales. The three Canadian sites record this abrupt shift between anno Domini 500 and 800, and subsequently conditions become increasingly variable. All three U.S. sites underwent a pronounced change, but the timing of this change is between anno Domini 1000 and 1300, thus later than in all of the Canadian sites. The mechanisms behind these patterns are poorly understood, but they are likely related to changes in the shape and location of the jet stream and associated storm tracks. If the patterns seen at these sites are representative of the region, this observed pattern can have huge implications for future water availability in this region. [ABSTRACT FROM AUTHOR]

The distribution and quality of water resources vary dramatically across Canada, and human impacts such as land-use and climate changes are exacerbating uncertainties in water supply and security. At the national level, Canada has no enforceable standards for safe drinking water and no comprehensive water-monitoring program to provide detailed, timely reporting on the state of water resources. To provide Canada's first national assessment of lake health, the NSERC Canadian Lake Pulse Network was launched in 2016 as an academic-government research partnership. LakePulse uses traditional approaches for limnological monitoring as well as state-of-the-art methods in the fields of genomics, emerging contaminants, greenhouse gases, invasive pathogens, paleolimnology, spatial modelling, statistical analysis, and remote sensing. A coordinated sampling program of about 680 lakes together with historical archives and a geomatics analysis of over 80,000 lake watersheds are used to examine the extent to which lakes are being altered now and in the future, and how this impacts aquatic ecosystem services of societal importance. Herein we review the network context, objectives and methods. Unlabelled Image • About 90% of Canada's municipal drinking water supplies are from surface waters. • An academic-government group is providing the first national review of lake health. • Over 100 variables were sampled at about 680 lakes across Canada. • Topics range from greenhouse gases to emerging contaminants to invasive pathogens. • The impacts of land-use and climate changes on lake health are also studied. [ABSTRACT FROM AUTHOR]

Calcium (Ca) is an essential element for almost all living organisms. Here, we examined global variation and controls of freshwater Ca concentrations, using 440 599 water samples from 43 184 inland water sites in 57 countries. We found that the global median Ca concentration was 4.0 mg L−1 with 20.7% of the water samples showing Ca concentrations ≤ 1.5 mg L−1, a threshold considered critical for the survival of many Ca-demanding organisms. Spatially, freshwater Ca concentrations were strongly and proportionally linked to carbonate alkalinity, with the highest Ca and carbonate alkalinity in waters with a pH around 8.0 and decreasing in concentrations towards lower pH. However, on a temporal scale, by analyzing decadal trends in >200 water bodies since the 1980s, we observed a frequent decoupling between carbonate alkalinity and Ca concentrations, which we attributed mainly to the influence of anthropogenic acid deposition. As acid deposition has been ameliorated, in many freshwaters carbonate alkalinity concentrations have increased or remained constant, while Ca concentrations have rapidly declined towards or even below pre-industrial conditions as a consequence of recovery from anthropogenic acidification. Thus, a paradoxical outcome of the successful remediation of acid deposition is a globally widespread freshwater Ca concentration decline towards critically low levels for many aquatic organisms. [ABSTRACT FROM AUTHOR]

Citizen science projects have a long history in ecological studies. The research usefulness of such projects is dependent on applying simple and standardized methods. Here, we conducted a citizen science project that involved more than 3500 Swedish high school students to examine the temperature difference between surface water and the overlying air (Tw-Ta) as a proxy for sensible heat flux (QH). If QH is directed upward, corresponding to positive Tw-Ta, it can enhance CO2 and CH4 emissions from inland waters, thereby contributing to increased greenhouse gas concentrations in the atmosphere. The students found mostly negative Tw-Ta across small ponds, lakes, streams/rivers and the sea shore (i.e. downward QH), with Tw-Ta becoming increasingly negative with increasing Ta. Further examination of Tw-Ta using high-frequency temperature data from inland waters across the globe confirmed that Tw-Ta is linearly related to Ta. Using the longest available high-frequency temperature time series from Lake Erken, Sweden, we found a rapid increase in the occasions of negative Tw-Ta with increasing annual mean Ta since 1989. From these results, we can expect that ongoing and projected global warming will result in increasingly negative Tw-Ta, thereby reducing CO2 and CH4 transfer velocities from inland waters into the atmosphere. [ABSTRACT FROM AUTHOR]

Calcium (Ca) concentrations are decreasing in softwater lakes across eastern North America and western Europe. Using long-term contemporary and palaeo-environmental field data, we show that this is precipitating a dramatic change in Canadian lakes: the replacement of previously dominant pelagic herbivores (Ca-rich Daphnia species) by Holopedium glacialis, a jelly-clad, Ca-poor competitor. In some lakes, this transformation is being facilitated by increases in macro-invertebrate predation, both from native (Chaoborus spp.) and introduced (Bythotrephes longimanus) zooplanktivores, to which Holopedium, with its jelly coat, is relatively invulnerable. Greater representation by Holopedium within cladoceran zooplankton communities will reduce nutrient transfer through food webs, given their lower phosphorus content relative to daphniids, and greater absolute abundances may pose long-term problems to water users. The dominance of jelly-clad zooplankton will likely persist while lakewater Ca levels remain low. [ABSTRACT FROM AUTHOR]

Although competing species are expected to exhibit compensatory dynamics (negative temporal covariation), empirical work has demonstrated that competitive communities often exhibit synchronous dynamics (positive temporal covariation). This has led to the suggestion that environmental forcing dominates species dynamics; however, synchronous and compensatory dynamics may appear at different length scales and/or at different times, making it challenging to identify their relative importance. We compiled 58 long-term datasets of zooplankton abundance in north-temperate and sub-tropical lakes and used wavelet analysis to quantify general patterns in the times and scales at which synchronous/compensatory dynamics dominated zooplankton communities in different regions and across the entire dataset. Synchronous dynamics were far more prevalent at all scales and times and were ubiquitous at the annual scale. Although we found compensatory dynamics in approximately 14% of all combinations of time period/scale/lake, there were no consistent scales or time periods during which compensatory dynamics were apparent across different regions. Our results suggest that the processes driving compensatory dynamics may be local in their extent, while those generating synchronous dynamics operate at much larger scales. This highlights an important gap in our understanding of the interaction between environmental and biotic forces that structure communities. [ABSTRACT FROM AUTHOR]