Indicators of the effects of climate change on freshwater ecosystems.

Freshwater ecosystems, including lakes, streams, and wetlands, are responsive to climate change and other natural and anthropogenic stresses. These ecosystems are frequently hydrologically and ecologically connected with one another and their surrounding landscapes, thereby integrating changes throughout their watersheds. The responses of any given freshwater ecosystem to climate change depend on the magnitude of climate forcing, interactions with other anthropogenic and natural changes, and the characteristics of the ecosystem itself. Therefore, the magnitude and manner in which freshwater ecosystems respond to climate change is difficult to predict a priori. We present a conceptual model to elucidate how freshwater ecosystems are altered by climate change. We identify eleven indicators that describe the response of freshwater ecosystems to climate change, discuss their potential value and limitations, and describe supporting measurements. Indicators are organized in three inter-related categories: hydrologic, water quality, and ecosystem structure and function. The indicators are supported by data sets with a wide range of temporal and spatial coverage, and they inform important scientific and management needs. Together, these indicators improve the understanding and management of the effects of climate change on freshwater ecosystems.

Identifying factors that affect mountain lake sensitivity to atmospheric nitrogen deposition across multiple scales.

Increased nitrogen (N) deposition rates over the past century have affected both North American and European mountain lake ecosystems. Ecological sensitivity of mountain lakes to N deposition varies, however, because chemical and biological responses are modulated by local watershed and lake properties. We evaluated predictors of mountain lake sensitivity to atmospheric N deposition across North American and European mountain ranges and included as response variables dissolved inorganic N (DIN = NNH4+ + NNO3-) concentrations and phytoplankton biomass. Predictors of these responses were evaluated at three different spatial scales (hemispheric, regional, subregional) using regression tree, random forest, and generalized additive model (GAM) analysis. Analyses agreed that Northern Hemisphere mountain lake DIN was related to N deposition rates and smaller scale spatial variability (e.g., regional variability between North American and European lakes, and subregional variability between mountain ranges). Analyses suggested that DIN, N deposition, and subregional variability were important for Northern Hemisphere mountain lake phytoplankton biomass. Together, these findings highlight the need for finer-scale, subregional analyses (by mountain range) of lake sensitivity to N deposition. Subregional analyses revealed differences in predictor variables of lake sensitivity. In addition to N deposition rates, lake and watershed features such as land cover, bedrock geology, maximum lake depth (Zmax), and elevation were common modulators of lake DIN. Subregional phytoplankton biomass was consistently positively related with total phosphorus (TP) in Europe, while North American locations showed variable relationships with N or P. This study reveals scale-dependent watershed and lake characteristics modulate mountain lake ecological responses to atmospheric N deposition and provides important context to inform empirically based management strategies.

Rock glaciers and related cold rocky landforms: Overlooked climate refugia for mountain biodiversity.

Mountains are global biodiversity hotspots where cold environments and their associated ecological communities are threatened by climate warming. Considerable research attention has been devoted to understanding the ecological effects of alpine glacier and snowfield recession. However, much less attention has been given to identifying climate refugia in mountain ecosystems where present-day environmental conditions will be maintained, at least in the near-term, as other habitats change. Around the world, montane communities of microbes, animals, and plants live on, adjacent to, and downstream of rock glaciers and related cold rocky landforms (CRL). These geomorphological features have been overlooked in the ecological literature despite being extremely common in mountain ranges worldwide with a propensity to support cold and stable habitats for aquatic and terrestrial biodiversity. CRLs are less responsive to atmospheric warming than alpine glaciers and snowfields due to the insulating nature and thermal inertia of their debris cover paired with their internal ventilation patterns. Thus, CRLs are likely to remain on the landscape after adjacent glaciers and snowfields have melted, thereby providing longer-term cold habitat for biodiversity living on and downstream of them. Here, we show that CRLs will likely act as key climate refugia for terrestrial and aquatic biodiversity in mountain ecosystems, offer guidelines for incorporating CRLs into conservation practices, and identify areas for future research.

Deeper waters are changing less consistently than surface waters in a global analysis of 102 lakes.

Globally, lake surface water temperatures have warmed rapidly relative to air temperatures, but changes in deepwater temperatures and vertical thermal structure are still largely unknown. We have compiled the most comprehensive data set to date of long-term (1970-2009) summertime vertical temperature profiles in lakes across the world to examine trends and drivers of whole-lake vertical thermal structure. We found significant increases in surface water temperatures across lakes at an average rate of + 0.37 °C decade-1, comparable to changes reported previously for other lakes, and similarly consistent trends of increasing water column stability (+ 0.08 kg m-3 decade-1). In contrast, however, deepwater temperature trends showed little change on average (+ 0.06 °C decade-1), but had high variability across lakes, with trends in individual lakes ranging from - 0.68 °C decade-1 to + 0.65 °C decade-1. The variability in deepwater temperature trends was not explained by trends in either surface water temperatures or thermal stability within lakes, and only 8.4% was explained by lake thermal region or local lake characteristics in a random forest analysis. These findings suggest that external drivers beyond our tested lake characteristics are important in explaining long-term trends in thermal structure, such as local to regional climate patterns or additional external anthropogenic influences.

Key rules of life and the fading cryosphere: Impacts in alpine lakes and streams.

Alpine regions are changing rapidly due to loss of snow and ice in response to ongoing climate change. While studies have documented ecological responses in alpine lakes and streams to these changes, our ability to predict such outcomes is limited. We propose that the application of fundamental rules of life can help develop necessary predictive frameworks. We focus on four key rules of life and their interactions: the temperature dependence of biotic processes from enzymes to evolution; the wavelength dependence of the effects of solar radiation on biological and ecological processes; the ramifications of the non-arbitrary elemental stoichiometry of life; and maximization of limiting resource use efficiency across scales. As the cryosphere melts and thaws, alpine lakes and streams will experience major changes in temperature regimes, absolute and relative inputs of solar radiation in ultraviolet and photosynthetically active radiation, and relative supplies of resources (e.g., carbon, nitrogen, and phosphorus), leading to nonlinear and interactive effects on particular biota, as well as on community and ecosystem properties. We propose that applying these key rules of life to cryosphere-influenced ecosystems will reduce uncertainties about the impacts of global change and help develop an integrated global view of rapidly changing alpine environments. However, doing so will require intensive interdisciplinary collaboration and international cooperation. More broadly, the alpine cryosphere is an example of a system where improving our understanding of mechanistic underpinnings of living systems might transform our ability to predict and mitigate the impacts of ongoing global change across the daunting scope of diversity in Earth's biota and environments.

Cross-ecosystem nutrient subsidies in Arctic and alpine lakes: implications of global change for remote lakes.

Environmental change is continuing to affect the flow of nutrients, material and organisms across ecosystem boundaries. These cross-system flows are termed ecosystem subsidies. Here, we synthesize current knowledge of cross-ecosystem nutrient subsidies between remote lakes and their surrounding terrain, cryosphere, and atmosphere. Remote Arctic and alpine lakes are ideal systems to study the effects of cross ecosystem subsidies because (a) they are positioned in locations experiencing rapid environmental changes, (b) they are ecologically sensitive to even small subsidy changes, (c) they have easily defined ecosystem boundaries, and (d) a variety of standard methods exist that allow for quantification of lake subsidies and their impacts on ecological communities and ecosystem functions. We highlight similarities and differences between Arctic and alpine systems and identify current knowledge gaps to be addressed with future work. It is important to understand the dynamics of nutrient and material flows between lakes and their environments in order to improve our ability to predict ecosystem responses to continued environmental change.

Environmental Controls on Microbial Diversity in Arctic Lakes of West Greenland.

We assessed the microbial community structure of six arctic lakes in West Greenland and investigated relationships to lake physical and chemical characteristics. Lakes from the ice sheet region exhibited the highest species richness, while inland and plateau lakes had lower observed taxonomical diversity. Lake habitat differentiation during summer stratification appeared to alter within lake microbial community composition in only a subset of lakes, while lake variability across regions was a consistent driver of microbial community composition in these arctic lakes. Principal coordinate analysis revealed differentiation of communities along two axes: each reflecting differences in morphometric (lake surface area), geographic (latitude and distance from the ice sheet), physical lake variables (water clarity), and lakewater chemistry (dissolved organic carbon [DOC], dissolved oxygen [DO], total nitrogen [TN], and conductivity). Understanding these relationships between environmental variables and microbial communities is especially important as heterotrophic microorganisms are key to organic matter decomposition, nutrient cycling, and carbon flow through nutrient poor aquatic environments in the Arctic.

Variable responses of dissolved organic carbon to precipitation events in boreal drinking water lakes.

In boreal regions, increased concentrations of dissolved organic carbon (DOC) have been linked to extreme wet years; however, less is known about the extent to which precipitation events are altering DOC concentration and quality. We assessed the effects of rain events on a suite of six lakes in Maine, U.S.A., to better understand how events alter DOC quantity and quality. DOC concentrations and DOC quality (measured as DOC-specific absorption coefficients (Specific Ultraviolet Absorbance (SUVA254 (also a*254), a*320, and a*380)) were quantified 24 h before, and at three time points (24-48 h, 5-7 days, and 3 weeks) after five different precipitation events. Our results revealed three types of responses across the lakes: (1) an initial spike in DOC concentrations of 30-133% and in the three quality metrics of 20-86% compared to pre-storm levels, followed by return to pre-storm concentrations; (2) a sustained increase in DOC concentrations (by 4-23%) and an increase in the three DOC quality metrics (by 1-43%) through the second post-storm sampling, with concentrations falling by the third post-storm sampling compared to pre-storm levels; and (3) no change during all sampling periods. Lake residence time was a key driver of changes in DOC concentration and DOC quality in response to storm events. Our research provides evidence that precipitation events contribute to short-term abrupt changes in DOC quantity and quality that are largely driven by key landscape and lake characteristics. These changes in DOC may have important implications for management of water utilities, including alteration or implementation of treatment strategies.

The Arctic in the Twenty-First Century: Changing Biogeochemical Linkages across a Paraglacial Landscape of Greenland.

The Kangerlussuaq area of southwest Greenland encompasses diverse ecological, geomorphic, and climate gradients that function over a range of spatial and temporal scales. Ecosystems range from the microbial communities on the ice sheet and moisture-stressed terrestrial vegetation (and their associated herbivores) to freshwater and oligosaline lakes. These ecosystems are linked by a dynamic glacio-fluvial-aeolian geomorphic system that transports water, geological material, organic carbon and nutrients from the glacier surface to adjacent terrestrial and aquatic systems. This paraglacial system is now subject to substantial change because of rapid regional warming since 2000. Here, we describe changes in the eco- and geomorphic systems at a range of timescales and explore rapid future change in the links that integrate these systems. We highlight the importance of cross-system subsidies at the landscape scale and, importantly, how these might change in the near future as the Arctic is expected to continue to warm.

Population divergence in fish elemental phenotypes associated with trophic phenotypes and lake trophic state.

Studies of ecological stoichiometry typically emphasize the role of interspecific variation in body elemental content and the effects of species or family identity. Recent work suggests substantial variation in body stoichiometry can also exist within species. The importance of this variation will depend on insights into its origins and consequences at various ecological scales, including the distribution of elemental phenotypes across landscapes and their role in nutrient recycling. We investigated whether trophic divergence can produce predictable patterns of elemental phenotypes among populations of an invasive fish, the white perch (Morone americana), and whether elemental phenotypes predict nutrient excretion. White perch populations exhibited a gradient of trophic phenotypes associated with landscape-scale variation in lake trophic state. Perch body chemistry varied considerably among lakes (from 0.09 for % C to 0.31-fold for % P) casting doubt on the assumption of homogenous elemental phenotypes. This variation was correlated with divergence in fish body shape and other trophic traits. Elemental phenotypes covaried (r (2) up to 0.84) with lake trophic state. This covariation likely arose in contemporary time since many of these perch populations were introduced in the last century and the trophic state in many of the lakes has changed in the past few decades. Nutrient excretion varied extensively among populations, but was not readily related to fish body chemistry or lake trophic state. This suggests that predictable patterns of fish body composition can arise quickly through trophic specialization to lake conditions, but such elemental phenotypes may not translate to altered nutrient recycling by fish.

Factors Controlling Methane in Arctic Lakes of Southwest Greenland.

We surveyed 15 lakes during the growing season of 2014 in Arctic lakes of southwest Greenland to determine which factors influence methane concentrations in these systems. Methane averaged 2.5 µmol L-1 in lakes, but varied a great deal across the landscape with lakes on older landscapes farther from the ice sheet margin having some of the highest values of methane reported in lakes in the northern hemisphere (125 µmol L-1). The most important factors influencing methane in Greenland lakes included ionic composition (SO4, Na, Cl) and chlorophyll a in the water column. DOC concentrations were also related to methane, but the short length of the study likely underestimated the influence and timing of DOC on methane concentrations in the region. Atmospheric methane concentrations are increasing globally, with freshwater ecosystems in northern latitudes continuing to serve as potentially large sources in the future. Much less is known about how freshwater lakes in Greenland fit in the global methane budget compared to other, more well-studied areas of the Arctic, hence our work provides essential data for a more complete view of this rapidly changing region.

Does allochthony in lakes change across an elevation gradient?.

Ecosystems are subsidized with inputs of mass and energy from their surroundings. These allochthonous inputs regulate many ecosystem characteristics. In inland waters, terrestrial inputs of organic matter regulate food-web structure, ecosystem metabolism, water clarity, and thermal stratification. Future changes in allochthony may be especially pronounced in high-elevation ecosystems due to increases in vegetation and precipitation associated with climate change. Several techniques exist to characterize the degree of allochthony of organic matter in aquatic systems, including metrics such as ΔH, the net isotopic discrimination between water and particulate organic matter (POM) of deuterium stable isotopes, and the fluorescence index (FI), which characterizes the fluorescence of dissolved organic matter (DOM). Despite the importance of allochthonous organic carbon inputs, little is known about either how allochthony varies across elevation gradients or whether different metrics are similarly related to allochthony. We measured AH, FI, and a suite of related water-quality characteristics in 30 lakes across a montane to alpine elevation gradient (2340 to 3205 m) in the Beartooth Mountains of Montana and Wyoming, USA, to understand how FI and AH varied with elevation, with one another, and with other allochthony-related water-quality characteristics. We hypothesized that allochthony of POM and DOM would decrease at higher elevations, with alpine lakes above treeline being more autochthonous compared with low-elevation lakes below treeline. We observed a significant inverse linear relationship between AH and Fl, with both metrics indicating a decrease in allochthony at higher elevations. Characteristics including the natural log of the ratio of concentrations of dissolved organic carbon to chlorophyll a (ln(DOC: Chl)), the spectral slope ratio between different spectra of two wavebands (SR, ratio of spectra at 275-295 to 350-400 nm), and a ratio of diffuse attenuation coefficients at 320 and 380 nm (KR, Kd320: Kd380) varied with both ΔH and FI while pH varied only with ΔH. High-elevation systems were characterized by low ln(DOC: Chl) and K(R), and high S(R) and pH. These results indicate that high-elevation lakes are more autochthonous than low-elevation lakes. The relationships among ΔH, FI, elevation, and other water-quality characteristics provide important insights to understand future changes in carbon cycling in mountain ecosystems.

Decadal trends reveal recent acceleration in the rate of recovery from acidification in the northeastern U.S.

Previous reports suggest variable trends in recovery from acidification in northeastern U.S. surface waters in response to the Clean Air Act Amendments. Here we analyze recent trends in emissions, wet deposition, and lake chemistry using long-term data from a variety of lakes in the Adirondack Mountains and New England. Sulfate concentration in wet deposition declined by more than 40% in the 2000s and sulfate concentration in lakes declined at a greater rate from 2002 to 2010 than during the 1980s or 1990s (-3.27 µeq L(-1)year(-1) as compared to -1.26 µeq L(-1)year(-1)). During the 2000s, nitrate concentration in wet deposition declined by more than 50% and nitrate concentration in lakes, which had no linear trend prior to 2000, declined at a rate of -0.05 µeq L(-1)year(-1). Base cation concentrations, which decreased during the 1990s (-1.5 µeq L(-1) year(-1)), have stabilized in New England lakes. Although total aluminum concentrations increased since 1999 (2.57 µg L(-1) year(-1)), there was a shift to nontoxic, organic aluminum. Despite this recent acceleration in recovery in multiple variables, both ANC and pH continue to have variable trends. This may be due in part to variable trajectories in the concentrations of base cations and dissolved organic carbon among our study lakes.

The influence of glacial meltwater on alpine aquatic ecosystems: a review.

The recent and rapid recession of alpine glaciers over the last 150 years has major implications for associated aquatic communities. Glacial meltwater shapes many of the physical features of high altitude lakes and streams, producing turbid environments with distinctive hydrology patterns relative to nival systems. Over the past decade, numerous studies have investigated the chemical and biological effects of glacial meltwater on freshwater ecosystems. Here, we review these studies across both lake and stream ecosystems. Focusing on alpine regions mainly in the Northern Hemisphere, we present examples of how glacial meltwater can affect habitat by altering physical and chemical features of aquatic ecosystems, and review the subsequent effects on the biological structure and function of lakes and streams. Collectively or separately, these factors can drive the overall distribution, diversity and behavior of primary producers, triggering cascading effects throughout the food web. We conclude by proposing areas for future research, particularly in regions where glaciers are soon projected to disappear.

Climate-induced changes in lake ecosystem structure inferred from coupled neo- and paleoecological approaches.

Over the 20th century, surface water temperatures have increased in many lake ecosystems around the world, but long-term trends in the vertical thermal structure of lakes remain unclear, despite the strong control that thermal stratification exerts on the biological response of lakes to climate change. Here we used both neo- and paleoecological approaches to develop a fossil-based inference model for lake mixing depths and thereby refine understanding of lake thermal structure change. We focused on three common planktonic diatom taxa, the distributions of which previous research suggests might be affected by mixing depth. Comparative lake surveys and growth rate experiments revealed that these species respond to lake thermal structure when nitrogen is sufficient, with species optima ranging from shallower to deeper mixing depths. The diatom-based mixing depth model was applied to sedimentary diatom profiles extending back to 1750 AD in two lakes with moderate nitrate concentrations but differing climate settings. Thermal reconstructions were consistent with expected changes, with shallower mixing depths inferred for an alpine lake where treeline has advanced, and deeper mixing depths inferred for a boreal lake where wind strength has increased. The inference model developed here provides a new tool to expand and refine understanding of climate-induced changes in lake ecosystems.

Mapping critical loads of nitrogen deposition for aquatic ecosystems in the Rocky Mountains, USA.

Spatially explicit estimates of critical loads of nitrogen (N) deposition (CL(Ndep)) for nutrient enrichment in aquatic ecosystems were developed for the Rocky Mountains, USA, using a geostatistical approach. The lowest CL(Ndep) estimates (<1.5 ± 1 kg N ha(-1) yr(-1)) occurred in high-elevation basins with steep slopes, sparse vegetation, and abundance of exposed bedrock and talus. These areas often correspond with areas of high N deposition (>3 kg N ha(-1) yr(-1)), resulting in CL(Ndep) exceedances ≥ 1.5 ± 1 kg N ha(-1) yr(-1). CL(Ndep) and CL(Ndep) exceedances exhibit substantial spatial variability related to basin characteristics and are highly sensitive to the NO(3)(-) threshold at which ecological effects are thought to occur. Based on an NO(3)(-) threshold of 0.5 µmol L(-1), N deposition exceeds CL(Ndep) in 21 ± 8% of the study area; thus, broad areas of the Rocky Mountains may be impacted by excess N deposition, with greatest impacts at high elevations.

A coherent signature of anthropogenic nitrogen deposition to remote watersheds of the Northern Hemisphere.

Humans have more than doubled the amount of reactive nitrogen (Nr) added to the biosphere, yet most of what is known about its accumulation and ecological effects is derived from studies of heavily populated regions. Nitrogen (N) stable isotope ratios ((15)N:(14)N) in dated sediments from 25 remote Northern Hemisphere lakes show a coherent signal of an isotopically distinct source of N to ecosystems beginning in 1895 ± 10 years (±1 standard deviation). Initial shifts in N isotope composition recorded in lake sediments coincide with anthropogenic CO(2) emissions but accelerate with widespread industrial Nr production during the past half century. Although current atmospheric Nr deposition rates in remote regions are relatively low, anthropogenic N has probably influenced watershed N budgets across the Northern Hemisphere for over a century.

Melting Alpine glaciers enrich high-elevation lakes with reactive nitrogen.

Alpine glaciers have receded substantially over the last century in many regions of the world. Resulting changes in glacial runoff not only affect the hydrological cycle, but can also alter the physical (i.e., turbidity from glacial flour) and biogeochemical properties of downstream ecosystems. Here we compare nutrient concentrations, transparency gradients, algal biomass, and fossil diatom species richness in two sets of high-elevation lakes: those fed by snowpack melt alone (SF lakes) and those fed by both glacial and snowpack meltwaters (GSF lakes). We found that nitrate (NO(3)(-)) concentrations in the GSF lakes were 1-2 orders of magnitude higher than in SF lakes. Although nitrogen (N) limitation is common in alpine lakes, algal biomass was lower in highly N-enriched GSF lakes than in the N-poor SF lakes. Contrary to expectations, GSF lakes were more transparent than SF lakes to ultraviolet and equally transparent to photosynthetically active radiation. Sediment diatom assemblages had lower taxonomic richness in the GSF lakes, a feature that has persisted over the last century. Our results demonstrate that the presence of glaciers on alpine watersheds more strongly influences NO(3)(-)concentrations in high-elevation lake ecosystems than any other geomorphic or biogeographic characteristic.

Quantifying recent ecological changes in remote lakes of North America and Greenland using sediment diatom assemblages.

BACKGROUND: Although arctic lakes have responded sensitively to 20(th)-century climate change, it remains uncertain how these ecological transformations compare with alpine and montane-boreal counterparts over the same interval. Furthermore, it is unclear to what degree other forcings, including atmospheric deposition of anthropogenic reactive nitrogen (Nr), have participated in recent regime shifts. Diatom-based paleolimnological syntheses offer an effective tool for retrospective assessments of past and ongoing changes in remote lake ecosystems. METHODOLOGY/PRINCIPAL FINDINGS: We synthesized 52 dated sediment diatom records from lakes in western North America and west Greenland, spanning broad latitudinal and altitudinal gradients, and representing alpine (n = 15), arctic (n = 20), and forested boreal-montane (n = 17) ecosystems. Diatom compositional turnover (beta-diversity) during the 20(th) century was estimated using Detrended Canonical Correspondence Analysis (DCCA) for each site and compared, for cores with sufficiently robust chronologies, to both the 19(th) century and the prior approximately 250 years (Little Ice Age). For both arctic and alpine lakes, beta-diversity during the 20(th) century is significantly greater than the previous 350 years, and increases with both latitude and altitude. Because no correlation is apparent between 20(th)-century diatom beta-diversity and any single physical or limnological parameter (including lake and catchment area, maximum depth, pH, conductivity, [NO(3)(-)], modeled Nr deposition, ambient summer and winter air temperatures, and modeled temperature trends 1948-2008), we used Principal Components Analysis (PCA) to summarize the amplitude of recent changes in relationship to lake pH, lake:catchment area ratio, modeled Nr deposition, and recent temperature trends. CONCLUSIONS/SIGNIFICANCE: The ecological responses of remote lakes to post-industrial environmental changes are complex. However, two regions reveal concentrations of sites with elevated 20(th)-century diatom beta-diversity: the Arctic where temperatures are increasing most rapidly, and mid-latitude alpine lakes impacted by high Nr deposition rates. We predict that remote lakes will continue to shift towards new ecological states in the Anthropocene, particularly in regions where these two forcings begin to intersect geographically.