Biodilution of Organic Species of Arsenic in Freshwater Food Webs.

Arsenic can accumulate in freshwater biota, sometimes reaching potentially harmful levels. However, the toxicity of arsenic strongly depends on which arsenic species are present. Although organic species are considered less harmful than inorganic ones, they have not been extensively studied in freshwater environments, and drivers of variation in arsenic speciation among sites and taxa remain unclear. We assessed concentrations of two organic arsenic species, arsenobetaine (AsB) and dimethylarsinic acid (DMA), in fish and invertebrates from three lakes near Sudbury, Ontario, Canada-a region with widespread mining impacts. Both AsB and DMA were detected in most samples (n = 212), varying across a wide range of concentrations (<0.001-30.144 and <0.006-5.262 mg/kg dry wt, respectively). The lake with the most severe mining impacts typically had the highest concentrations (designated by square brackets []) of AsB and DMA. In contrast, the percentage of total arsenic made up by AsB (%AsB) and DMA (%DMA) did not vary significantly between lakes. Arsenic speciation in fish muscle varied with fish size, selenium concentrations, and trophic elevation (inferred from nitrogen stable isotope ratios δ15N), but relationships with dietary carbon source (inferred from carbon stable isotope ratios δ13C) were more varied. Within all three lake food webs, [AsB] and [DMA] typically underwent biodilution, decreasing with trophic elevation (i.e., δ15N). Although the aforementioned factors explained some variation in arsenic speciation, there remains considerable unexplained variation. Further studies on arsenic speciation in freshwater biota should target a wider diversity of taxa to better understand drivers of variation in arsenic speciation. In addition, research emphasizing the percentage of inorganic arsenic and other organic arsenic species is needed to improve environmental and human health risk assessments. Environ Toxicol Chem 2024;43:833-846. © 2024 The Authors. Environmental Toxicology and Chemistry published by Wiley Periodicals LLC on behalf of SETAC.

The speciation of arsenic in the muscle tissue of inland and coastal freshwater fish from a remote boreal region.

Elevated concentrations of total arsenic (As) have been reported in boreal freshwater fish in both human-impacted and relatively pristine areas. We assessed the arsenic speciation profiles in muscle tissue of six fish species (n = 300) sampled from nine locations across a remote freshwater watershed in northern Ontario, Canada, extending from inland headwater lakes to the coastal marine confluence. Of the five arsenic species measured, only arsenobetaine (AsB) and dimethylarsinic acid (DMA) were detected in these fish. Riverine fish had up to 10-fold higher total [As] when compared to lacustrine fish. On average, these riverine fish also had higher percentages of AsB (%AsB, 60 ± 26%) and lower percentages of unmeasured arsenic (%UNM, 20 ± 21%), compared to lacustrine fish (28 ± 18% and 52 ± 21% %AsB and %UNM, respectively). DMA percentages (%DMA) were relatively consistent across the watershed, averaging 20 ± 21% across all fish. We examined ecological drivers of As speciation and found that %AsB increased slightly with fish weight in large-body predatory fish, but not in forage fish or insectivores. Furthermore, %AsB was positively related to trophic elevation (inferred from δ15N) in lacustrine fish across 3 out of 4 communities and within some populations. Lastly, riverine fish with a more marine-based diet had markedly higher %AsB when compared to fish with more freshwater-based diets, indicating an effect of anadromy on arsenic speciation. Overall, knowledge on arsenic speciation in freshwater fish has been limited and these results indicate potential drivers that can be considered in future studies. Furthermore, the absence of toxic inorganic As species in these boreal fish is an important consideration for future environmental monitoring practices and risk assessments, some of which assume 10-20% of total [As] in fish is present as toxic inorganic As. Additional studies on As bioaccumulation and biotransformation are needed in freshwater systems, particularly at the base of aquatic food webs.

Age, body size, growth and dietary habits: What are the key factors driving individual variability in mercury of lacustrine fishes in northern temperate lakes?

Fish total mercury concentration ([THg]) has been linked to various fish attributes, but the relative importance of these attributes in accounting for among-individual variation in [THg] has not been thoroughly assessed. We compared the contributions of ontogeny (age, length), growth (growth rate, body condition), and food web position (δ13C, δ15N) to among-individual variability in [THg] within populations of seven common fishes from 141 north temperate lakes. Ontogenetic factors accounted for most variation in [THg]; age was a stronger and less variable predictor than length for most species. Adding both indices of growth and food web position to these models increased explained variation (R2) in [THg] by 6-25% among species. Fish [THg] at age increased with growth rate, while fish [THg] at length decreased with growth rate, and the effect of body condition was consistently negative. Trophic elevation (inferred from δ15N) was a stronger predictor than primary production source (inferred from δ13C) for piscivores but not benthivores. Fish [THg] increased with δ15N in all species but showed a more variable relationship with δ13C. Among-individual variation in [THg] is primarily related to age or size in most temperate freshwater fishes, and effects of growth rate and food web position need to be considered in the context of these ontogenetic drivers.

Forest defoliator outbreaks alter nutrient cycling in northern waters.

Insect defoliators alter biogeochemical cycles from land into receiving waters by consuming terrestrial biomass and releasing biolabile frass. Here, we related insect outbreaks to water chemistry across 12 boreal lake catchments over 32-years. We report, on average, 27% lower dissolved organic carbon (DOC) and 112% higher dissolved inorganic nitrogen (DIN) concentrations in lake waters when defoliators covered entire catchments and reduced leaf area. DOC reductions reached 32% when deciduous stands dominated. Within-year changes in DOC from insect outbreaks exceeded 86% of between-year trends across a larger dataset of 266 boreal and north temperate lakes from 1990 to 2016. Similarly, within-year increases in DIN from insect outbreaks exceeded local, between-year changes in DIN by 12-times, on average. As insect defoliator outbreaks occur at least every 5 years across a wider 439,661 km2 boreal ecozone of Ontario, we suggest they are an underappreciated driver of biogeochemical cycles in forest catchments of this region.

Modifying effects of a cobble substrate on thermal environments and implications for embryonic development in lake whitefish (Coregonus clupeaformis).

A laboratory flume was constructed to examine substrate effects on aquatic development. The flume was designed as a once-through system with a submerged cobble-filled corebox. Lake whitefish (Coregonus clupeaformis) embryos and temperature probes were deployed at multiple sites within the cobble and in the open water channel. Embryos were incubated in the flume for two different experimental periods: one to examine substrate impacts during natural lake cooling (37 days: 5 December 2016 to 10 January 2017) and the second to investigate substrate effects while administering a twice weekly 1 h heat shock (51 days: 11 January to 2 March 2017). During incubation, no significant difference was found in the average temperature between locations; however, temperatures were more stable within the cobble. Following both incubation periods, embryos retrieved from the cobble were significantly smaller in both dry mass and body length by up to 20%. These results demonstrate differences between embryos submerged in a cobble substrate and in the open water column, highlighting the need to consider the physical influences from the incubation environment when assessing development effects as part of any scientific study or environmental assessment.

Arsenic, chromium, and other elements of concern in fish from remote boreal lakes and rivers: Drivers of variation and implications for subsistence consumption.

Eating fish provides numerous health benefits, but it is also a dominant pathway for human exposure to contaminants. Many studies have examined mercury (Hg) accumulation in fish, but fewer have considered other elements, such as arsenic (As) and chromium (Cr). Recently, freshwater fish from several pristine boreal systems across northern Ontario, Canada, have been reported with elevated concentrations of As and Cr for reasons that are not well understood. Our goal was to investigate the ecological and environmental influences over concentrations of As, Cr, and other elements in these fish to better understand what affects metal uptake and the risk to consumers. We measured 10 elements (including As, Cr, Hg) as well as carbon (δ13C), nitrogen (δ15N), and sulfur (δ34S) stable isotopes in 388 fish from 25 lake and river sites across this remote region. These data were used to determine the effect of: 1) trophic ecology; and 2) watershed geology on piscine elemental content. Overall, most element concentrations were low, often below provincial advisory benchmarks (ABs). However, traces of Hg, As, Cr, and selenium (Se) were detected in most fish. Based on their exceedance of their respective ABs, the most restrictive elements on fish consumption in these boreal systems were Hg > As > Cr. Arsenic and Se, but not Cr concentrations were related to fish size and trophic ecology (inferred from δ13C and δ15N), suggesting bioaccumulation of the former elements. Fish with enriched δ34S values, suggestive of anadromous behaviour, had marginally lower Hg but higher Se concentrations. Modeling results suggested a strong effect of site-specific factors, though we found weak trends between piscine elemental content and geological features (e.g., mafic intrusions), potentially due to the broad spatial scale of this study. Results from this study address gaps in our understanding of As and Cr bioaccumulation and will help to inform fish consumption guidelines.

Microbiome functioning depends on individual and interactive effects of the environment and community structure.

How ecosystem functioning changes with microbial communities remains an open question in natural ecosystems. Both present-day environmental conditions and historical events, such as past differences in dispersal, can have a greater influence over ecosystem function than the diversity or abundance of both taxa and genes. Here, we estimated how individual and interactive effects of microbial community structure defined by diversity and abundance, present-day environmental conditions, and an indicator of historical legacies influenced ecosystem functioning in lake sediments. We studied sediments because they have strong gradients in all three of these ecosystem properties and deliver important functions worldwide. By characterizing bacterial community composition and functional traits at eight sites fed by discrete and contrasting catchments, we found that taxonomic diversity and the normalized abundance of oxidase-encoding genes explained as much variation in CO2 production as present-day gradients of pH and organic matter quantity and quality. Functional gene diversity was not linked to CO2 production rates. Surprisingly, the effects of taxonomic diversity and normalized oxidase abundance in the model predicting CO2 production were attributable to site-level differences in bacterial communities unrelated to the present-day environment, suggesting that colonization history rather than habitat-based filtering indirectly influenced ecosystem functioning. Our findings add to limited evidence that biodiversity and gene abundance explain patterns of microbiome functioning in nature. Yet we highlight among the first time how these relationships depend directly on present-day environmental conditions and indirectly on historical legacies, and so need to be contextualized with these other ecosystem properties.

Percentage of methylmercury in the muscle tissue of freshwater fish varies with body size and age and among species.

It is commonly assumed that most (>95%) of the mercury (Hg) found in fish muscle is the toxic form, methylmercury (MeHg), due to its efficient assimilation and retention in biotic tissue. However, this assumption is largely based on studies examining the percentage of MeHg (%MeHg [the fraction of total Hg as MeHg]) in muscle from mostly large-bodied predatory fish; less is known about the %MeHg in smaller bodied individuals or those of different trophic guilds. The present study analyzed MeHg and total Hg concentrations in the muscle of 2 large-bodied piscivores (walleye and northern pike), one large-bodied benthivore (white sucker), and 2 small-bodied forage fish (sculpins and shiners) across a broad size range. We found substantially lower %MeHg than the commonly assumed 95% in several fish (e.g., 17 individuals had <70% MeHg). Muscle %MeHg significantly increased with size and age in all species except walleye, which had significantly higher %MeHg than pike or suckers, particularly in smaller and younger fish (e.g., 18-21% higher at 10 g; 5-11% higher at 500 g). Results of predictive modeling suggest that muscle %MeHg is higher in pelagic-feeding fish and those with lower lipid content, although model results varied significantly among species. According to our findings, total Hg measurement in muscle is not an appropriate proxy for MeHg in smaller fish from all species, an important consideration for future piscine Hg studies and monitoring. Environ Toxicol Chem 2018;37:2682-2691. © 2018 SETAC.

Optical Properties of Dissolved Organic Matter and Their Relation to Mercury Concentrations in Water and Biota Across a Remote Freshwater Drainage Basin.

Dissolved organic matter (DOM) includes an array of carbon-based compounds that vary in size and structure and have complex interactions with mercury (Hg) cycling in aquatic systems. While many studies have examined the relationship between dissolved organic carbon concentrations ([DOC]) and methyl Hg bioaccumulation, few studies have considered the effects of DOM composition (e.g., protein-content, aromaticity). The goal of this study was to explore the relationships between total and methyl [Hg] in water, invertebrates, and fish and optically derived measures of DOM composition from 47 lake and river sites across a boreal watershed. Results showed higher aqueous total [Hg] in systems with more aromatic DOM and higher [DOC], potentially due to enhanced transport from upstream or riparian areas. Methyl [Hg] in biota were all positively related to the amount of microbial-based DOM and, in some cases, to the proportions of labile and protein-like DOM. These results suggest that increased Hg bioaccumulation is related to the availability of labile DOM, potentially due to enhanced Hg methylation. DOM composition explained 68% and 54% more variability in [Hg] in surface waters and large-bodied fish, respectively, than [DOC] alone. These results show that optical measures of DOM characteristics are a valuable tool for understanding DOM-Hg biogeochemistry.

Subcatchment deltas and upland features influence multiscale aquatic ecosystem recovery in damaged landscapes.

Assessing biological recovery in damaged aquatic environments requires the consideration of multiple spatial and temporal scales. Past research has focused on assessing lake recovery from atmospheric or catchment disturbance at regional or catchment levels. Studies have also rarely considered the influences of adjacent terrestrial characteristics on within-lake habitats, such as subcatchment delta confluences. We used Hyalella azteca, a ubiquitous freshwater amphipod, as a sensitive indicator to assess the importance of local subcatchment scale factors in the context of multiscale lake recovery within the metal mining region of Sudbury, Canada following a period of major reductions in atmospheric pollution. At the regional scale, data from repeated surveys of 40 lakes showed higher probabilities of H. azteca occurrence with higher lake water conductivity, alkalinity, and pH and lower metal concentrations. The importance of metals decreased through time and the importance of higher conductivity, alkalinity, and pH increased. At the subcatchment scale, a subset of six lakes sampled across a colonization gradient revealed higher H. azteca abundances at subcatchment delta sites than non-delta sites in early colonization stages, and that abundance at delta sites was correlated with both within-lake habitat and terrestrial subcatchment characteristics. For example, wetland cover reduced the strength of positive associations between H. azteca abundance and macrophyte density. A single lake from this subset also revealed higher abundances at delta sites associated with higher concentrations of terrestrial organic matter and larger subcatchments. Our results demonstrate that factors affecting recovery can change with the scale of study, and that managing terrestrial-aquatic linkages is important for facilitating recovery processes within damaged lake ecosystems.

Terrestrial support of lake food webs: Synthesis reveals controls over cross-ecosystem resource use.

Widespread evidence that organic matter exported from terrestrial into aquatic ecosystems supports recipient food webs remains controversial. A pressing question is not only whether high terrestrial support is possible but also what the general conditions are under which it arises. We assemble the largest data set, to date, of the isotopic composition (δ2H, δ13C, and δ15N) of lake zooplankton and the resources at the base of their associated food webs. In total, our data set spans 559 observations across 147 lakes from the boreal to subtropics. By predicting terrestrial resource support from within-lake and catchment-level characteristics, we found that half of all consumer observations that is, the median were composed of at least 42% terrestrially derived material. In general, terrestrial support of zooplankton was greatest in lakes with large physical and hydrological connections to catchments that were rich in aboveground and belowground organic matter. However, some consumers responded less strongly to terrestrial resources where within-lake production was elevated. Our study shows that multiple mechanisms drive widespread cross-ecosystem support of aquatic consumers across Northern Hemisphere lakes and suggests that changes in terrestrial landscapes will influence ecosystem processes well beyond their boundaries.

Dry conditions disrupt terrestrial-aquatic linkages in northern catchments.

Aquatic ecosystems depend on terrestrial organic matter (tOM) to regulate many functions, such as food web production and water quality, but an increasing frequency and intensity of drought across northern ecosystems is threatening to disrupt this important connection. Dry conditions reduce tOM export and can also oxidize wetland soils and release stored contaminants into stream flow after rainfall. Here, we test whether these disruptions to terrestrial-aquatic linkages occur during mild summer drought and whether this affects biota across 43 littoral zone sites in 11 lakes. We use copper (Cu) and nickel (Ni) as representative contaminants, and measure abundances of Hyalella azteca, a widespread indicator of ecosystem condition and food web production. We found that tOM concentrations were reduced but correlations with organic soils (wetlands and riparian forests) persisted during mild drought and were sufficient to suppress labile Cu concentrations. Wetlands, however, also became a source of labile Ni to littoral zones, which was linked to reduced abundances of the amphipod H. azteca, on average by up to 70 times across the range of observed Ni concentrations. This reveals a duality in the functional linkage of organic soils to aquatic ecosystems whereby they can help buffer the effects of hydrologic disconnection between catchments and lakes but at the cost of biogeochemical changes that release stored contaminants. As evidence of the toxicity of trace contaminant concentrations and their global dispersion grows, sustaining links among forests, organic soils and aquatic ecosystems in a changing climate will become increasingly important.

Predicting chronic copper and nickel reproductive toxicity to Daphnia pulex-pulicaria from whole-animal metabolic profiles.

The emergence of omics approaches in environmental research has enhanced our understanding of the mechanisms underlying toxicity; however, extrapolation from molecular effects to whole-organism and population level outcomes remains a considerable challenge. Using environmentally relevant, sublethal, concentrations of two metals (Cu and Ni), both singly and in binary mixtures, we integrated data from traditional chronic, partial life-cycle toxicity testing and metabolomics to generate a statistical model that was predictive of reproductive impairment in a Daphnia pulex-pulicaria hybrid that was isolated from an historically metal-stressed lake. Furthermore, we determined that the metabolic profiles of organisms exposed in a separate acute assay were also predictive of impaired reproduction following metal exposure. Thus we were able to directly associate molecular profiles to a key population response - reproduction, a key step towards improving environmental risk assessment and management.

Metabolomics confirms that dissolved organic carbon mitigates copper toxicity.

Reductions in atmospheric emissions from the metal smelters in Sudbury, Canada, produced major improvements in acid and metal contamination of local lakes and indirectly increased dissolved organic carbon (DOC) concentrations. Metal toxicity, however, has remained a persistent problem for aquatic biota. Integrating high-throughput, nontargeted mass spectrometry metabolomics with conventional toxicological measures elucidated the mediating effects of dissolved organic matter (DOM) on the toxicity of Cu to Daphnia pulex-pulicaria, a hybrid isolated from these soft water lakes. Two generations of daphniids were exposed to Cu (0-20 µg/L) at increasing levels of natural DOM (0-4 mg DOC/L). Added DOM reduced Cu toxicity monotonically with median lethal concentration values increasing from 2.3 µg/L Cu without DOM to 22.7 µg/L Cu at 4 mg DOC/L. Reproductive output similarly benefited, increasing with DOM, yet falling with increases in Cu. Second generation reproduction was more impaired than the first generation. Dissolved organic matter had a greater influence than Cu on the metabolic status of the daphniids. Putative identification of metabolite peaks indicated that DOM elevation increased the metabolic energy status of the first generation animals, but this benefit was reduced in the second generation, although evidence of increased oxidative stress was detected. These results indicate that Sudbury's terrestrial ecosystems should be managed to increase aquatic DOM supply to enable daphniid colonists to both survive and foster stable populations.

Forests fuel fish growth in freshwater deltas.

Aquatic ecosystems are fuelled by biogeochemical inputs from surrounding lands and within-lake primary production. Disturbances that change these inputs may affect how aquatic ecosystems function and deliver services vital to humans. Here we test, using a forest cover gradient across eight separate catchments, whether disturbances that remove terrestrial biomass lower organic matter inputs into freshwater lakes, thereby reducing food web productivity. We focus on deltas formed at the stream-lake interface where terrestrial-derived particulate material is deposited. We find that organic matter export increases from more forested catchments, enhancing bacterial biomass. This transfers energy upwards through communities of heavier zooplankton, leading to a fourfold increase in weights of planktivorous young-of-the-year fish. At least 34% of fish biomass is supported by terrestrial primary production, increasing to 66% with greater forest cover. Habitat tracers confirm fish were closely associated with individual catchments, demonstrating that watershed protection and restoration increase biomass in critical life-stages of fish.

Temporal changes in mercury concentrations of large-bodied fishes in the boreal shield ecoregion of northern Ontario, Canada.

Much of the mercury (Hg) in freshwater fish of the boreal shield ecoregion is believed to originate from atmospheric deposition. As such, declines in fish Hg concentrations would be expected in response to recent declines in atmospheric Hg deposition in this ecoregion. We compared recent (2005-2010) and historic (1974-1981) muscle total mercury concentrations ([THg], standardized to a fish body mass of 1 kg) in seven fish species (five piscivores, two benthivores) from 73 lakes in northern Ontario (Canada) using a paired-comparisons approach. The rate of bioaccumulation (i.e., slopes of log(e)[THg] vs log(e) total length relationship) increased for walleye (Sander vitreus) but did not change significantly for any other species. There was no significant decline in mean [THg] between recent and historic time periods for any species. In fact, recent mean [THg] were slightly higher (<0.08 ppm) than historic mean [THg] for all species, and this difference was significant for northern pike (Esox lucius). The magnitude of the temporal change in northern pike declined significantly from south to north over the study area but there were no discernible geographic patterns in the temporal change in [THg] for any other species. This study shows that [THg] of most large-bodied fish species in boreal shield lakes are not declining in response to the decline in atmospheric Hg deposition.

Assessing disturbance from under-ice noise on fishes in boreal lakes.

There are many potential sources of anthropogenic noise that can manifest under ice in boreal lakes that are within the hearing ranges of northern boreal fishes. Impacts of noise on fish can be correlated to the fish's hearing sensitivity. In general, boreal fishes have most sensitive hearing <400 Hz,but this varies by species and life stage. By assessing the hearing capabilities and vocalizations of fish species and sound signatures from development activities, it may be possible to forecast potential impacts resulting from expected under-ice noise exposure.There is still the need for basic information to facilitate the assessment and identification of potential impacts (e.g., the hearing capabilities of many species of boreal fishes at different life stages and the sound signatures of various anthropogenic noise sources). Additionally, the impacts to fish from a particular noise source (e.g., pile driving) or the potential for a sound-related disturbance ata particular time in a species life history, such as courtship or spawning, warrants further study.

Recovery of benthic invertebrate communities from acidification in Killarney Park lakes.

Using a reference-condition comparison, recovery of benthic invertebrate communities from acidification was assessed in three lakes in Killarney Wilderness Park approximately 40-60 km from the massive metal smelters in Sudbury, Canada. Test site analyses (TSAs) were used to compare the park lakes to 20 reference lakes near Dorset Ontario, 200 km to the east. An extension of a previous survey (1997-2001) of two sensitive mayfly species (Stenonema femoratum and Stenacron interpunctatum) was conducted in one of the lakes. TSA results indicate that the three Killarney lakes remain significantly different from reference condition due primarily to higher abundances of a few acid-tolerant families and the presence of some less abundant sensitive families. Colonization rates differ greatly between the two mayfly species presumably because of competition for available habitat. Overall, this study suggests that early colonizers will gain an advantage to out-compete subsequent arrivals, and these competitive interactions will delay the return of communities to reference condition.

Sediment trace metal profiles in lakes of Killarney Park, Canada: from regional to continental influence.

The lakes in Killarney Provincial Park (KPP) located 40-60 km southwest of Sudbury, Ontario are beginning to recover after decades of being severely affected by acidification and atmospheric pollutants. Detailed profiles of acid-recoverable trace elements (As. Cd, Cu, Co. Fe, Mn, Ni, Pb and Zn) were obtained after aqua regia digestion and ICP-OES analysis of sediment cores taken from six Park lakes. Results permitted the identification of two types of profiles. The first type applies to elements such as Fe, Mn, As and Co for which historical deposition and recent recovery are strongly masked by diagenetic remobilization. The second type of profile applies to elements such as Cd, Cu, Ni, Pb and Zn on which the history of industrialisation in North America and mining activities in Sudbury can be superimposed. Based on sediment data of trace elements less affected by diagenetic remobilization (Cd, Cu, Ni, Pb, Zn), chemical recovery indices can be estimated from depth profiles. Indices of maximum (C(p)) and surface (C(s)) contamination were calculated by dividing the concentration of a given metal by the pre-industrial level. The ratio of the two indices provided a simple estimation of the chemical recovery of lakes that does not consider the influence of the watershed or the lake pH. Profiles of metals in sediment of KPP complement the water quality monitoring data and tend to indicate that this area is in transition from dominant influence of regional pollution sources to becoming controlled by continental atmospheric deposition.

Developing conceptual frameworks for the recovery of aquatic biota from acidification.

Surface water acidity is decreasing in large areas of Europe and North America in response to reductions in atmospheric S deposition, but the ecological responses to these water-quality improvements are uncertain. Biota are recovering in some lakes and rivers, as water quality improves, but they are not yet recovering in others. To make sense of these different responses, and to foster effective management of the acid rain problem, we need to understand 2 things: i) the sequence of ecological steps needed for biotic communities to recover; and ii) where and how to intervene in this process should recovery stall. Here our purpose is to develop conceptual frameworks to serve these 2 needs. In the first framework, the primarily ecological one, a decision tree highlights the sequence of processes necessary for ecological recovery, linking them with management tools and responses to bottlenecks in the process. These bottlenecks are inadequate water quality, an inadequate supply of colonists to permit establishment, and community-level impediments to recovery dynamics. A second, more management-oriented framework identifies where we can intervene to overcome these bottlenecks, and what research is needed to build the models to operationalize the framework. Our ability to assess the benefits of S emission reduction would be simplified if we had models to predict the rate and extent of ecological recovery from acidification. To build such models we must identify the ecological steps in the recovery process. The frameworks we present will advance us towards this goal.