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Stormwater ponds are common features in urbanized landscapes because they enhance flood reduction and nutrient retention. With shallow depths and high inputs of organic matter, these systems can be highly productive with rapid oxygen depletion when thermally stratified or ice‐covered. However, most of our understanding of the biogeochemistry of stormwater ponds comes from the open water period. We explored under‐ice oxygen dynamics in 20 stormwater ponds in Madison, WI (USA) that were ice covered from late December to early March to investigate the drivers of bottom water oxygen saturation and the impact on the accumulation of carbon dioxide (CO2) and methane (CH4). Winter anoxia was driven by ice transmissivity, winter nutrient concentrations, and precedent summer productivity. Oxygen depletion led to overall higher concentrations of greenhouse gases in pond surface waters. This research enhances our understanding of winter pond biogeochemistry and its links to summer productivity. Plain Language Summary: In urban areas, stormwater ponds are common to help manage flooding and filter out nutrients to improve water quality in downstream water bodies. In the summer, urban ponds can experience a loss of oxygen in the bottom water, but less is known about what happens in these ponds in the winter under ice cover. We measured oxygen profiles in 20 ponds during the winter to investigate the drivers of oxygen loss in the bottom waters. We also explored the consequence of oxygen loss by measuring greenhouse gas (carbon dioxide and methane) concentrations in the surface water. Winter oxygen loss was driven by snow and ice clarity, winter nutrients, and previous summer organic matter from either algae or floating plants. Ponds with less oxygen under ice had higher concentrations of greenhouse gases in the surface water. Warmer winters could increase freeze thaw cycles and decrease oxygen under ice or increase oxygen with more instances of open water. Key Points: Urban ponds exhibit a range of under‐ice oxygen regimesWinter anoxia is driven by ice transmissivity, nutrient concentrations, and preceding summer productivityA consequence of under‐ice oxygen depletion is the accumulation of methane and carbon dioxide [ABSTRACT FROM AUTHOR]

Microcystin poses a serious threat to aquatic ecosystems and human health. There is a pressing need to understand the production, movement, and storage of microcystin in lakes. We constructed a conceptual biogeochemical model for microcystin through a comprehensive literature synthesis, identifying four major pools and nine major fluxes in lakes that also connect to the terrestrial environment. This conceptual model can be used as the framework for developing ecosystem mass balances of microcystin. We propose that the concentration of microcystin in the water column is the balance between the import, sediment translocation, production and degradation, uptake, burial, and export. However, substantial unknowns remain pertaining to the magnitude and movement of microcystin. Future investigations should focus on sediment fluxes, drivers of biodegradation, and seasonal dynamics. Adopting the framework of a “microcystin cycle” improves our understanding of processes driving toxin prevalence and helps to prioritize strategies for minimizing exposure risks. [ABSTRACT FROM AUTHOR]

Martin A. Simonson, Michael J. Weber, Grace M. Wilkinson and Andrew R. Annear. 2023. Annual changes in water quality and sportfish community structure following commercial harvest of common carp and bigmouth buffalo. Lake Reserv Manage. 39:174–189. Commercial harvest of common carp (Cyprinus carpio; hereafter carp) and bigmouth buffalo (Ictiobus cyprinellus; hereafter buffalo) populations had little detectable effect on shallow lake ecosystems. We tested whether carp and buffalo biomass removal affects limnological variables and fish community metrics across 6 shallow, natural lakes of northwestern Iowa using mixed effects models. Annual commercial harvest of carp ranged from 0 to 71 kg/ha; annual harvest of buffalo ranged from 0 to 356 kg/ha. Biomanipulation (i.e., commercial harvest) of carp was associated with decreases in soluble phosphorus and nitrogen concentrations, but not total phosphorus or nitrogen. Buffalo harvest was unrelated to annual changes in nutrient concentrations but was associated with reductions in chlorophyll a and phycocyanin concentrations. Secchi disk transparency and total suspended solids were unrelated to carp and buffalo harvest. Carp harvest was associated with reduced biomass of large cladocerans but no other zooplankton biomass densities; buffalo harvest was unrelated to zooplankton biomass. Species richness and rake density of aquatic macrophytes were unrelated to carp and buffalo harvest. Carp and buffalo harvest was unrelated to changes in most indices of sportfish abundance, condition, and size distribution. Our results suggest harvest of carp and buffalo <71 kg/ha has little effect on abiotic and biotic ecosystem components on short time scales and highlights the challenges associated with shallow lake restoration. [ABSTRACT FROM AUTHOR]

Phytoplankton blooms often follow nutrient enrichment. Differences among lakes in light‐absorbing dissolved organic carbon (DOC) may shift bloom thresholds to higher nutrient loads and thereby increase resilience of lakes to enrichment. To explore this idea, we measured resilience to experimental enrichment of two lakes with contrasting DOC concentrations. We compared bloom thresholds in both lakes using a model of phytoplankton response to DOC and nutrients, a dynamic time series indicator of resilience, and two empirical measures of stochastic resilience, mean exit time and median survival time. For the dynamic indicator and ecosystem model the lake with higher DOC was more resilient to enrichment. However, the distributions overlapped for stochastic indicators of resilience of the two lakes. These analyses show that DOC interacts with mixing depth and zooplankton biomass to affect resilience. Strong contrasts in DOC and many observations are needed to discern effects of DOC on resilience to enrichment. [ABSTRACT FROM AUTHOR]

Consumer nutrient recycling influences aquatic ecosystem functioning by altering the movement and transformation of nutrients. In hypereutrophic reservoirs, zooplankton nutrient recycling has been considered negligible due to high concentrations of available nutrients. A comparative analysis (Moody and Wilkinson, 2019) found that zooplankton communities in hypereutrophic lakes are dominated by nitrogen (N)-rich species, which the authors hypothesized would increase phosphorus (P) availability through excretion. However, zooplankton nutrient recycling likely varies over the course of a growing season due to changes in biomass, community composition and grazing pressure on phytoplankton. We quantified zooplankton, phytoplankton and nutrient concentration dynamics during the summer of 2019 in a temperate, hypereutrophic reservoir. We found that the estimated contribution of zooplankton excretion to the dissolved nutrient pool on a given day was equivalent to a substantial proportion (21–39%) of the dissolved inorganic P standing stock in early summer when P concentrations were low and limiting phytoplankton growth. Further, we found evidence that zooplankton affected phytoplankton size distributions through selective grazing of smaller phytoplankton cells likely affecting nutrient uptake and storage by phytoplankton. Overall, our results demonstrate zooplankton excretion in hypereutrophic reservoirs likely helped drive springtime phytoplankton dynamics through nutrient recycling while grazing influenced phytoplankton size distributions. [ABSTRACT FROM AUTHOR]

Phosphorus (P) flux across the sediment–water interface in lakes and reservoirs responds to external perturbations within the context of sediment characteristics. Lentic ecosystems experience profound spatiotemporal heterogeneity in the mechanisms that control sediment P fluxes, likely producing hot spots and hot moments of internal loading. However, spatiotemporal variation in P fluxes remains poorly quantified, particularly in the context of sediment chemistry as a controlling variable. We measured P flux rates and mobile sediment P forms along the longitudinal gradient of a temperate reservoir every 2 months from February to October 2020. Both aerobic and anaerobic processes mobilized sediment P throughout the year. High flux rates at littoral sampling sites (8.4 and 9.7 mg P m−2 day−1) occurred in late summer under oxic conditions in the overlying water and mobilized labile organic P. High fluxes at the profundal site coincided with hypolimnetic anoxia under ice cover and in mid‐summer (11.2 and 17.2 mg P m−2 day−1, respectively) and released redox‐sensitive P. Several high fluxes substantially skewed the flux rate distribution, providing evidence of hot spots and hot moments of internal loading. We further scaled the measured sediment P flux rates to representative areas of the lakebed to estimate internal P loads at an ecosystem scale. We found that P release from littoral sites under oxic conditions in the overlying water had an outsized impact on total loads. Our findings demonstrate the importance of considering spatial and seasonal variation in sediment P pools and fluxes in order to more accurately estimate internal loads and identify the dominant biogeochemical mechanisms involved. [ABSTRACT FROM AUTHOR]

Phosphorus (P) release from lakebed sediments may fuel phytoplankton blooms, especially in shallow waterbodies. A primary mechanism that controls internal P loading is the size and chemical composition of the sediment P pool. However, variation in sediment P within and among shallow lakes remains poorly quantified. We measured the degree of spatial heterogeneity in the size and composition of sediment P pools, both within and among seven shallow lakes. There was a 1.6× difference in total sediment P among the study lakes, and P composition varied across lakes based on differences in watershed soils and land cover and lake basin morphology. Differences in sediment P composition among lakes imply that the dominant mechanisms of internal loading will also vary among lakes. We also found that higher mobile P (as a fraction of total P) in the profundal sediments was positively correlated with long‐term mean chlorophyll‐a concentrations (p = 0.04), indicating the reciprocal relationship between sediment P composition and phytoplankton biomass in shallow lakes. Additionally, we measured substantial within‐lake heterogeneity in total and loosely‐bound sediment P within each lake. Concentrations were positively correlated with water depth such that extrapolating measurements from the deep site alone could overestimate whole‐lake mean P concentrations, reinforcing that single station methods produce unreliable estimates of mean sediment P stocks. Our results provide insight into the magnitude and pattern of inter‐ and intra‐lake variation in sediment P pools that should be accounted for when sampling, scaling measurements, and modeling sediment P dynamics. Plain Language Summary: Phosphorus (P) is an essential nutrient in freshwater ecosystems; however, in excess it can cause blooms of algae and cyanobacteria. P enters lakes from the watershed (external loading) or when released from lakebed sediments into the overlying water (internal loading). Sediment P chemistry controls internal loading because some forms of P are more susceptible to release. However, we lack a quantitative understanding of how sediment P chemistry can vary within individual lakes and among different waterbodies. We measured variation in sediment P composition within and among seven shallow lakes. We found that sediment P composition varied across lakes based on lake and watershed characteristics. We also found that a higher abundance of mobile P forms in the sediments was correlated with higher concentrations of phytoplankton pigments, indicating that sediment P composition may influence phytoplankton biomass in shallow lakes. Within individual study lakes, sediment P was highly variable, meaning that multiple samples are required to capture this heterogeneity. Our results reveal the scale of spatial variation in sediment P forms in shallow lakes. Since sediment P composition is a primary mechanism controlling sediment P release, this knowledge is critical to building accurate models to predict internal P loading. Key Points: The size and chemical composition of sediment phosphorus pools are spatially heterogeneous both within and among shallow lakesLake basin morphology, water depth, and watershed characteristics drive variation in sediment P pools within and among lakesUnderstanding intra‐lake variation in sediment P informs the spatial sampling resolutions necessary for accurate P stock assessments [ABSTRACT FROM AUTHOR]

Algal blooms, the rapid proliferation of algal biomass often to nuisance or harmful levels, diminish aquatic ecosystem services. Freshwater blooms can cause substantial economic damage by interrupting water supply, limiting recreation, and reducing property values. The interaction between eutrophication and climate change has been hypothesized to drive widespread intensification of blooms in inland waters, although there is little empirical evidence that this trend is pervasive. Here, we show that bloom intensification in inland waterbodies – defined as trends in chlorophyll‐a of increasing bloom magnitude, severity, or duration – has not been widespread for hundreds of lakes in the US. Only 10.8% of the 323 waterbodies analyzed had significant bloom intensification. Conversely, 16.4% of the waterbodies had significant decreasing trends during the same period. While it is encouraging that bloom intensification is not currently widespread, continued efforts toward aquatic ecosystem protection and restoration are imperative for maintaining ecosystem services into the future. [ABSTRACT FROM AUTHOR]

Microevolution can have consequences at higher levels of ecological organization. Although divergence among populations can be rapid and driven by anthropogenic changes to the environment, the ecological relevance of evolution induced by human activities remains poorly understood.A frequent way in which human activities drive microevolution is the increase in supply of nutrients such as phosphorus (P) that are required for fitness‐relevant traits such as growth and reproduction. Because higher P concentrations decrease P‐use efficiency and feeding rate in heterotrophic consumers such as Daphnia, we hypothesized that such adjustments should alter consumer–resource dynamics.We examined how cultural eutrophication in temperate lakes causes trait variation in the grazer Daphnia pulicaria. We tested for variation in Daphnia traits and genetic variation in the metabolic enzyme phosphoglucose isomerase (Pgi) which are each known to respond to eutrophication. We then examined the impact of this variation on consumer–resource dynamics using a combination of experiments and a multi‐lake survey.We found that Daphnia from hypereutrophic lakes responded to experimental hypereutrophic conditions with increased growth rates and fecundity when raised on P‐fertilized seston, but had reduced performance on P‐poor seston relative to eutrophic source Daphnia. These results suggest that Daphnia may face a trade‐off in performance at low versus excess P that may be mediated in part by genetic variation at the Pgi locus.The variation observed in laboratory growth experiments scaled up to Daphnia populations in both mesocosm experiments and among lakes. In both the mesocosm experiment and in the lake survey, Daphnia from hypereutrophic source lakes reached high biomass while phytoplankton biomass also remained high.Given the prevalence and rapid eutrophication of freshwater ecosystems worldwide, these results indicate that considering the potential effects of evolutionary change in ecosystem models could be useful in forecasting the effects of anthropogenic environmental change on pivotal ecosystem services. [ABSTRACT FROM AUTHOR]

Algal blooms can have profound effects on the structure and function of aquatic ecosystems and have the potential to interrupt valuable ecosystem services. Despite the potential ecological and economic consequences of algal blooms, the spatial dynamics of bloom development in spatially complex ecosystems such as shallow lakes remain poorly characterised. Our goal was to evaluate the magnitude and drivers of spatial variability of algal biomass, dissolved oxygen, and pH over the course of a season, in a shallow lake in order to better understand the spatial dynamics of algal blooms in these ecosystems.We sampled 98 locations in a small eutrophic lake on a 65‐m grid for several parameters (chlorophyll a, phycocyanin, dissolved oxygen, pH, and temperature), weekly over 122 days. This was done to estimate the dynamics of variability and spatial autocorrelation during the course of multiple bloom events. We also compared the spatial measurements to a high frequency sensor deployed at a fixed station and estimated the optimal spatial sampling resolution by performing a rarefaction analysis.Spatial heterogeneity of algal pigments was high, particularly during bloom events, and this pattern and the overall severity of the bloom were not well captured with the fixed station monitoring. The pattern of algal pigments and other limnologically important variables (dissolved oxygen and pH) was related to the direction of prevailing winds 24 hr prior to sampling, the shallow northern basin where the main surface inlet is located, and heavy precipitation. Additionally, a dense bed of floating‐leaf macrophytes contributed to local patchiness in all variables. Finally, from the rarefaction analysis we found that minimal information about the mean state of the ecosystem was gained after c. 30 locations had been sampled.This study revealed how spatially heterogeneous shallow lakes are over the course of a single season, and that the magnitude of variability was highest during biologically intensive periods such as algal blooms. As such, continued research is needed across a range of trophic conditions to better understand the structure of horizontal variability in lakes. Overall, these data demonstrate the need for spatially explicit monitoring to better understand the dynamics and drivers of algal blooms in shallow lakes and to better manage ecosystem services. [ABSTRACT FROM AUTHOR]

The effects that fishes have on rates of one or more ecosystem processes (hereafter referred to as functional effects) are often invoked as an important reason for imperilled species conservation. However, the degree to which we understand these effects is rarely evaluated for most groups of fishes. We assessed how well the functional effects of freshwater and diadromous fishes, one of the most imperilled groups of animals, are quantified to date. We found that 88% of studies considering the functional effects of imperilled North American fishes were conducted on one family, Salmonidae. Studies of the functional effects of fishes were also concentrated in Pacific drainages of North America, with few studies in hotspots of imperilled fish diversity such as the Southeastern United States, the arid Southwest and central Mexico. Our results demonstrate the vast taxonomic and geographic gaps in our functional understanding of imperilled fishes and highlight the need to broaden this work to justify the argument that they are functionally important in the ecosystems they inhabit. [ABSTRACT FROM AUTHOR]

Functional variation among consumer communities can alter ecosystem nutrient cycling. These impacts on ecosystem function can be specifically driven by interspecific variation in stoichiometric traits; thus, functional trait‐based approaches can be used to explain the processes controlling ecosystem stoichiometry. However, eutrophication may reduce the functional importance of consumers in ecosystems by eliminating heterogeneity in nutrient recycling among taxa.To test whether zooplankton functional diversity, i.e. aspects of the stoichiometric trait space occupied by zooplankton communities, varies over gradients in trophic state and nutrient stoichiometry, we examined functional and taxonomic variation in the zooplankton communities of 130 lakes in the agriculturally dominated state of Iowa (U.S.A.) over 7 years.Stoichiometric functional dispersion decreased with trophic state index, supporting the trait abundance shift hypothesis that hypereutrophic lakes are characterised by different combinations of functional traits than their less eutrophic counterparts. Zooplankton communities became increasingly N‐rich relative to P as TSI increased. Specifically, P‐poor Bosmina, Chydorus, and cyclopoid copepods increased in abundance with eutrophication.Stoichiometric trait distributions of zooplankton shift with eutrophication, which implies that the unique functioning of hypereutrophic lakes could be due in part to the consumers inhabiting them. As zooplankton N:P increased with trophic state while lake total nitrogen to total phosphorus ratio decreased with trophic state, P‐poor zooplankton taxa may exacerbate excess P availability in these hypereutrophic systems by differentially recycling P at higher rates. [ABSTRACT FROM AUTHOR]

Terrestrial organic matter can be assimilated by aquatic consumers but implications for biomass and production are unresolved. An ecosystem model was fit to estimate effects of phosphorus (P) load, planktivory, and supply rate of terrestrial particulate organic carbon ( TPOC) on phytoplankton and zooplankton in five whole-lake experiments. Phytoplankton biomass increased with P load and planktivory and decreased with TPOC supply rate. Zooplankton biomass increased with P load and responded weakly to planktivory and TPOC supply rate. Zooplankton allochthony (proportion of carbon from terrestrial sources) decreased with P load and planktivory and increased with TPOC supply rate. Lakes with low allochthony (< 0.3) had wide ranges of phytoplankton and zooplankton biomass and production, depending on P load and planktivory. Lakes with high allochthony (> 0.3) had low biomass and production of both phytoplankton and zooplankton. In summary, terrestrial OC inhibits primary production and is a relatively low-quality food source for zooplankton. [ABSTRACT FROM AUTHOR]

Hydrogen stable isotopes ( δ2H) are used for quantifying resources supporting food webs. However, application of δ2H in mixing models requires; (1) correction for environmental water ( ω) in consumer tissues, (2) consideration of differential fractionation among biochemical constituents, and (3) consideration of differential H-exchange among samples and standards. We present data and sensitivity analyses addressing each of these issues and provide recommendations for future isotope food web studies. First, we determined from field data that maximum ω for aquatic consumers averaged 0.23 ± 0.03, similar to the median ω from a survey of published values (0.22 ± 0.02). Resource use estimates based solely on δ2H data were sensitive to the selected ω value. Second, to quantify the potential bias in bulk tissue analysis from differential tissue fractionation, we calculated the change in whole organism δ2H before and after lipid extraction for 61 aquatic samples. The average change in consumers' δ2H after lipid extraction was a positive shift of 11.8‰ relative to the pre-extraction value. This shift resulted in a minor change in resource use estimates when correcting for lipids. Finally, we evaluated the impact of correcting for H-exchange in samples using standards with dissimilar H-exchange portions. The impact of the correction factor for H-exchange on resource use estimates could be large if suitable standards are not used for comparison. From these analyses we conclude that despite these complicating factors, analysis of resource use is possible using whole organisms' δ2H, especially in combination with cautionary sensitivity analysis. [ABSTRACT FROM AUTHOR]

Lake food webs can be supported by primary production from within the lake, organic matter imported from the catchment or some mixture of these two sources. Generalisations about food-web subsidies to lake ecosystems are often based on data from only a few ecosystems and therefore do not consider the potential variability of subsidies among diverse ecosystems in a landscape., We measured the variation among lake ecosystems in terrestrial (allochthonous) utilisation by pelagic consumers and developed models to describe the variability. Stable isotope ratios of hydrogen and carbon were measured for Chaoborus spp. and crustacean zooplankton taxa in 40 lakes to quantify consumer allochthonous resource use (allochthony)., The median fraction of consumer allochthony estimated using a two-source Bayesian mixing model varied between 4 and 82% (mean among all lakes = 36%) for Chaoborus sp. among lakes and between 1 and 76% in a more limited sample of crustacean zooplankton consumers. The degree of allochthonous resource use increased linearly with the availability of allochthonous resources., Terrestrial support of Chaoborus was correlated (using best fitting relationships) with covariates for lake organic matter sources including dissolved inorganic carbon, total phosphorus, chlorophyll α, colour and catchment area. However, the most parsimonious model was an inverse relationship between lake surface area and consumer allochthony, indicating that allochthonous subsidies are more important in smaller than larger systems. Given the preponderance of small waterbodies, allochthonous subsidies are important in a large number of lake ecosystems. [ABSTRACT FROM AUTHOR]

Aquatic ecosystems are hotspots of decomposition and sources of carbon dioxide to the atmosphere that are globally significant. Carbon exported from land (allochthonous) also supplements the carbon fixed by photosynthesis in aquatic ecosystems (autochthonous), contributing to the organic matter (OM) that supports aquatic consumers. Although the presence of terrestrial compounds in aquatic OM is well known, the contribution of terrestrial versus aquatic sources to the composition of OM has been quantified for only a handful of systems. Here we use stable isotopes of hydrogen and carbon to demonstrate that the terrestrial contribution (ΦTerr) to particulate organic matter (POM) is as large or larger (mean = 54.6% terrestrial) than the algal contribution in 39 lakes of the northern highlands region of Wisconsin and Michigan. Further, the largest carbon pool, dissolved organic matter (DOM), is strongly dominated by allochthonous material (mean for the same set of lakes approximately 100% terrestrial). Among lakes, increases in terrestrial contribution to POM are significantly correlated with more acidic pH. Extrapolating this relationship using a survey of pH in 1692 lakes in the region reveals that, with the exception of eutrophic lakes, most of the OM in lakes is of terrestrial origin. These results are consistent with the growing evidence that lakes are significant conduits for returning degraded terrestrial carbon to the atmosphere. [ABSTRACT FROM AUTHOR]

Resources in lakes are vertically partitioned due to stratification and trophic interactions. Metalimnetic phytoplankton could be an important resource for zooplankton that either reside in the metalimnion or migrate through this layer. However, it is difficult to estimate metalimnetic resource use, especially using isotope approaches, because surface and deep phytoplankton often have similar isotopic compositions. To overcome this limitation, we experimentally enriched the metalimnetic dissolved inorganic carbon (DIC) 13C pool in Peter Lake to enhance the isotopic separation between metalimnetic phytoplankton and other resources. Metalimnetic δ13C-DIC peaked at 73.2‰ after the isotope addition and maintained an average enrichment of 34.5‰ above epilimnetic δ13C-DIC for 62 d. Combining hydrogen and carbon stable isotope values, we estimated the epilimnetic, metalimnetic, and terrestrial resource use by zooplankton, using a Bayesian mixing model that accounted for uncertainties in both consumers and sources. We also measured diel vertical migration and net ecosystem production with in situ bottle incubations over the course of the experiment. Metalimnetic resource use was minor (0-8%) for zooplankton that either resided in the epilimnion of the lake during the day or migrated there at night. For consumers that resided in the metalimnion, metalimnetic phytoplankton accounted for 18-21% of zooplankton isotope composition. The most important resource for all zooplankton was terrestrial organic matter (56-73% of consumer mass), regardless of habitat. This experiment indicates that, in lakes like Peter Lake, metalimnetic autochthonous resources are of minor importance to zooplankton relative to epilimnetic autochthonous and allochthonous resources. [ABSTRACT FROM AUTHOR]

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 (δ 2 H, δ 13 C, and δ 15 N) 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.