Global patterns of allochthony in stream-riparian meta-ecosystems.

Ecosystems that are coupled by reciprocal flows of energy and nutrient subsidies can be viewed as a single "meta-ecosystem." Despite these connections, the reciprocal flow of subsidies is greatly asymmetrical and seasonally pulsed. Here, we synthesize existing literature on stream-riparian meta-ecosystems to quantify global patterns of the amount of subsidy consumption by organisms, known as "allochthony." These resource flows are important since they can comprise a large portion of consumer diets, but can be disrupted by human modification of streams and riparian zones. Despite asymmetrical subsidy flows, we found stream and riparian consumer allochthony to be equivalent. Although both fish and stream invertebrates rely on seasonally pulsed allochthonous resources, we find allochthony varies seasonally only for fish, being nearly three times greater during the summer and fall than during the winter and spring. We also find that consumer allochthony varies with feeding traits for aquatic invertebrates, fish, and terrestrial arthropods, but not for terrestrial vertebrates. Finally, we find that allochthony varies by climate for aquatic invertebrates, being nearly twice as great in arid climates than in tropical climates, but not for fish. These findings are critical to understanding the consequences of global change, as ecosystem connections are being increasingly disrupted.

Assessing placement bias of the global river gauge network.

Knowing where and when rivers flow is paramount to managing freshwater ecosystems. Yet stream gauging stations are distributed sparsely across rivers globally and may not capture the diversity of fluvial network properties and anthropogenic influences. Here we evaluate the placement bias of a global stream gauge dataset on its representation of socioecological, hydrologic, climatic and physiographic diversity of rivers. We find that gauges are located disproportionally in large, perennial rivers draining more human-occupied watersheds. Gauges are sparsely distributed in protected areas and rivers characterized by non-perennial flow regimes, both of which are critical to freshwater conservation and water security concerns. Disparities between the geography of the global gauging network and the broad diversity of streams and rivers weakens our ability to understand critical hydrologic processes and make informed water-management and policy decisions. Our findings underscore the need to address current gauge placement biases by investing in and prioritizing the installation of new gauging stations, embracing alternative water-monitoring strategies, advancing innovation in hydrologic modelling, and increasing accessibility of local and regional gauging data to support human responses to water challenges, both today and in the future.

White-tailed deer consumption of emergent macrophytes mediates aquatic-to-terrestrial nutrient flows.

Trophic interactions between mobile animals and their food sources often vector resource flows across ecosystem boundaries. However, the quality and quantity of such ecological subsidies may be altered by indirect interactions between seemingly unconnected taxa. We studied whether emergent macrophytes growing at the aquatic-terrestrial interface facilitate multi-step aquatic-to-terrestrial resource flows between streams and terrestrial herbivores. We also explored whether aquatic animal aggregations indirectly promote such resource flows by creating biogeochemical hotspots of nutrient cycling and availability.We tested whether white-tailed deer (Odocoileus virginianus) in eastern North America vector nutrient fluxes from streams to terrestrial ecosystems by consuming emergent macrophytes (Justicia americana) using isotope and nutrient analyses of fecal samples and motion-sensing cameras. We also tested whether mussel-generated biogeochemical hotspots might promote such fluxes by surveying the density and nutrient stoichiometry of J. americana beds growing in association with variable densities of freshwater mussels (Bivalvia: Unionoida).Fecal samples from riparian deer had 3% lower C:N and 20% lower C:P ratios than those in upland habitats. C and N isotopes suggested riparian deer ate both terrestrial and aquatic (J. americana) vegetation, whereas upland deer ate more terrestrial foods. Motion-sensing cameras showed deer eating J. americana more than twice as frequently at mussel-generated hotspots than non-mussel sites. However, mussels were not associated with variation in J. americana growth or N and P content-although N isotopes in J. americana leaves did suggest assimilation of animal-derived nutrients.Our findings suggest that white-tailed deer may conduct significant transfers of aquatic-derived nutrients into terrestrial habitats when they feed on macrophytes and defecate on land. Whether aquatic animal aggregations promote such resource flows by creating biogeochemical hotspots remains unresolved, but the nearly global distributions of the deer family (Cervidae) and of macrophytes suggest that cervid-driven aquatic-to-terrestrial nutrient flows may be widespread and ecologically important.

Reconceptualizing the hyporheic zone for nonperennial rivers and streams.

Nonperennial streams dominate global river networks and are increasing in occurrence across space and time. When surface flow ceases or the surface water dries, flow or moisture can be retained in the subsurface sediments of the hyporheic zone, supporting aquatic communities and ecosystem processes. However, hydrological and ecological definitions of the hyporheic zone have been developed in perennial rivers and emphasize the mixing of water and organisms, respectively, from both the surface stream and groundwater. The adaptation of such definitions to include both humid and dry unsaturated conditions could promote characterization of how hydrological and biogeochemical variability shape ecological communities within nonperennial hyporheic zones, advancing our understanding of both ecosystem structure and function in these habitats. To conceptualize hyporheic zones for nonperennial streams, we review how water sources and surface and subsurface structure influence hydrological and physicochemical conditions. We consider the extent of this zone and how biogeochemistry and ecology might vary with surface states. We then link these components to the composition of nonperennial stream communities. Next, we examine literature to identify priorities for hydrological and ecological research exploring nonperennial hyporheic zones. Lastly, by integrating hydrology, biogeochemistry, and ecology, we recommend a multidisciplinary conceptualization of the nonperennial hyporheic zone as the porous subsurface streambed sediments that shift between lotic, lentic, humid, and dry conditions in space and time to support aquatic-terrestrial biodiversity. As river drying increases in extent because of global change, we call for holistic, interdisciplinary research across the terrestrial and aquatic sciences to apply this conceptualization to characterize hyporheic zone structure and function across the full spectrum of hydrological states.

A global synthesis of human impacts on the multifunctionality of streams and rivers.

Human impacts, particularly nutrient pollution and land-use change, have caused significant declines in the quality and quantity of freshwater resources. Most global assessments have concentrated on species diversity and composition, but effects on the multifunctionality of streams and rivers remain unclear. Here, we analyse the most comprehensive compilation of stream ecosystem functions to date to provide an overview of the responses of nutrient uptake, leaf litter decomposition, ecosystem productivity, and food web complexity to six globally pervasive human stressors. We show that human stressors inhibited ecosystem functioning for most stressor-function pairs. Nitrate uptake efficiency was most affected and was inhibited by 347% due to agriculture. However, concomitant negative and positive effects were common even within a given stressor-function pair. Some part of this variability in effect direction could be explained by the structural heterogeneity of the landscape and latitudinal position of the streams. Ranking human stressors by their absolute effects on ecosystem multifunctionality revealed significant effects for all studied stressors, with wastewater effluents (194%), agriculture (148%), and urban land use (137%) having the strongest effects. Our results demonstrate that we are at risk of losing the functional backbone of streams and rivers if human stressors persist in contemporary intensity, and that freshwaters are losing critical ecosystem services that humans rely on. We advocate for more studies on the effects of multiple stressors on ecosystem multifunctionality to improve the functional understanding of human impacts. Finally, freshwater management must shift its focus toward an ecological function-based approach and needs to develop strategies for maintaining or restoring ecosystem functioning of streams and rivers.

Cadmium-Induced Kidney Injury: Oxidative Damage as a Unifying Mechanism.

Cadmium is a nonessential metal that has heavily polluted the environment due to human activities. It can be absorbed into the human body via the gastrointestinal tract, respiratory tract, and the skin, and can cause chronic damage to the kidneys. The main site where cadmium accumulates and causes damage within the nephrons is the proximal tubule. This accumulation can induce dysfunction of the mitochondrial electron transport chain, leading to electron leakage and production of reactive oxygen species (ROS). Cadmium may also impair the function of NADPH oxidase, resulting in another source of ROS. These ROS together can cause oxidative damage to DNA, proteins, and lipids, triggering epithelial cell death and a decline in kidney function. In this article, we also reviewed evidence that the antioxidant power of plant extracts, herbal medicines, and pharmacological agents could ameliorate cadmium-induced kidney injury. Finally, a model of cadmium-induced kidney injury, centering on the notion that oxidative damage is a unifying mechanism of cadmium renal toxicity, is also presented. Given that cadmium exposure is inevitable, further studies using animal models are warranted for a detailed understanding of the mechanism underlying cadmium induced ROS production, and for the identification of more therapeutic targets.

Energy pathways modulate the resilience of stream invertebrate communities to drought.

While climate change is altering ecosystems on a global scale, not all ecosystems are responding in the same way. The resilience of ecological communities may depend on whether food webs are producer- or detritus-based (i.e. 'green' or 'brown' food webs, respectively), or both (i.e. 'multi-channel' food web). Food web theory suggests that the presence of multiple energy pathways can enhance community stability and resilience and may modulate the responses of ecological communities to disturbances such as climate change. Despite important advances in food web theory, few studies have empirically investigated the resilience of ecological communities to climate change stressors in ecosystems with different primary energy channels. We conducted a factorial experiment using outdoor stream mesocosms to investigate the independent and interactive effects of warming and drought on invertebrate communities in food webs with different energy channel configurations. Warming had little effect on invertebrates, but stream drying negatively impacted total invertebrate abundance, biomass, richness and diversity. Although resistance to drying did not differ among energy channel treatments, recovery and overall resilience were higher in green mesocosms than in mixed and brown mesocosms. Resilience to drying also varied widely among taxa, with larger predatory taxa exhibiting lower resilience. Our results suggest that the effects of drought on stream communities may vary regionally and depend on whether food webs are fuelled by autochthonous or allochthonous basal resources. Communities inhabiting streams with large amounts of organic matter and more complex substrates that provide refugia may be more resilient to the loss of surface water than communities inhabiting streams with simpler, more homogeneous substrates.

River ecosystem conceptual models and non-perennial rivers: A critical review.

Conceptual models underpin river ecosystem research. However, current models focus on continuously flowing rivers and few explicitly address characteristics such as flow cessation and drying. The applicability of existing conceptual models to nonperennial rivers that cease to flow (intermittent rivers and ephemeral streams, IRES) has not been evaluated. We reviewed 18 models, finding that they collectively describe main drivers of biogeochemical and ecological patterns and processes longitudinally (upstream-downstream), laterally (channel-riparian-floodplain), vertically (surface water-groundwater), and temporally across local and landscape scales. However, perennial rivers are longitudinally continuous while IRES are longitudinally discontinuous. Whereas perennial rivers have bidirectional lateral connections between aquatic and terrestrial ecosystems, in IRES, this connection is unidirectional for much of the time, from terrestrial-to-aquatic only. Vertical connectivity between surface and subsurface water occurs bidirectionally and is temporally consistent in perennial rivers. However, in IRES, this exchange is temporally variable, and can become unidirectional during drying or rewetting phases. Finally, drying adds another dimension of flow variation to be considered across temporal and spatial scales in IRES, much as flooding is considered as a temporally and spatially dynamic process in perennial rivers. Here, we focus on ways in which existing models could be modified to accommodate drying as a fundamental process that can alter these patterns and processes across spatial and temporal dimensions in streams. This perspective is needed to support river science and management in our era of rapid global change, including increasing duration, frequency, and occurrence of drying.

What's in a Name? Patterns, Trends, and Suggestions for Defining Non-Perennial Rivers and Streams.

Rivers that cease to flow are globally prevalent. Although many epithets have been used for these rivers, a consensus on terminology has not yet been reached. Doing so would facilitate a marked increase in interdisciplinary interest as well as critical need for clear regulations. Here we reviewed literature from Web of Science database searches of 12 epithets to learn (Objective 1-O1) if epithet topics are consistent across Web of Science categories using latent Dirichlet allocation topic modeling. We also analyzed publication rates and topics over time to (O2) assess changes in epithet use. We compiled literature definitions to (O3) identify how epithets have been delineated and, lastly, suggest universal terms and definitions. We found a lack of consensus in epithet use between and among various fields. We also found that epithet usage has changed over time, as research focus has shifted from description to modeling. We conclude that multiple epithets are redundant. We offer specific definitions for three epithets (non-perennial, intermittent, and ephemeral) to guide consensus on epithet use. Limiting the number of epithets used in non-perennial river research can facilitate more effective communication among research fields and provide clear guidelines for writing regulatory documents.

Zero or not? Causes and consequences of zero-flow stream gage readings.

Streamflow observations can be used to understand, predict, and contextualize hydrologic, ecological, and biogeochemical processes and conditions in streams. Stream gages are point measurements along rivers where streamflow is measured, and are often used to infer upstream watershed-scale processes. When stream gages read zero, this may indicate that the stream has fully dried; however, zero-flow readings can also be caused by a wide range of other factors. Our ability to identify whether or not a zero-flow gage reading indicates a dry fluvial system has far reaching environmental implications. Incorrect identification and interpretation by the data user can lead to hydrologic, ecological, and/or biogeochemical predictions from models and analyses. Here, we describe several causes of zero-flow gage readings: frozen surface water, flow reversals, instrument error, and natural or human-driven upstream source losses or bypass flow. For these examples, we discuss the implications of zero-flow interpretations. We also highlight additional methodss for determining flow presence, including direct observations, statistical methods, and hydrologic models, which can be applied to interpret causes of zero-flow gage readings and implications for reach- and watershed-scale dynamics. Such efforts are necessary to improve our ability to understand and predict surface flow activation, cessation, and connectivity across river networks. Developing this integrated understanding of the wide range of possible meanings of zero-flows will only attain greater importance in a more variable and changing hydrologic climate.

Animal aggregations promote emergent aquatic plant production at the aquatic-terrestrial interface.

The roles mobile animals and abiotic processes play as vectors for resource transfers between ecosystems ("subsidies") are well studied, but the idea that resources from animals with limited mobility may be transported across boundaries through intermediate taxa remains unexplored. Aquatic plants ("macrophytes") are globally distributed and may mediate transfers of aquatic-derived nutrients from aggregations of aquatic animals to terrestrial ecosystems when consumed by terrestrial herbivores. We used mesocosms (94 × 44 cm) to test whether aquatic animal-generated biogeochemical hotspots increase growth and nutrient content in macrophytes using the macrophyte Justicia americana and freshwater mussels. Justicia americana biomass production increased and belowground biomass allocation changed with increasing mussel density. At high mussel density, water-column phosphorus increased and carbon:phosphorus ratios in J. americana tissues decreased. We deployed motion-sensing cameras to explore herbivory on J. americana growing along the margins of the Kiamichi River, Oklahoma, and documented feeding by large mammals (Odocoileus virginianus, Sus scrofa, and Bos taurus). Thus, biogeochemical hotspots generated by aquatic animal aggregations can promote macrophyte production that subsequently is transferred to terrestrial animals. More broadly, this suggests that reductions in aquatic animal biomass may have bottom-up impacts that indirectly affect terrestrial ecosystems via plant-animal interactions bridging ecosystem boundaries.

Global patterns and drivers of ecosystem functioning in rivers and riparian zones.

River ecosystems receive and process vast quantities of terrestrial organic carbon, the fate of which depends strongly on microbial activity. Variation in and controls of processing rates, however, are poorly characterized at the global scale. In response, we used a peer-sourced research network and a highly standardized carbon processing assay to conduct a global-scale field experiment in greater than 1000 river and riparian sites. We found that Earth's biomes have distinct carbon processing signatures. Slow processing is evident across latitudes, whereas rapid rates are restricted to lower latitudes. Both the mean rate and variability decline with latitude, suggesting temperature constraints toward the poles and greater roles for other environmental drivers (e.g., nutrient loading) toward the equator. These results and data set the stage for unprecedented "next-generation biomonitoring" by establishing baselines to help quantify environmental impacts to the functioning of ecosystems at a global scale.

Dynamical implications of bi-directional resource exchange within a meta-ecosystem.

The exchange of resources across ecosystem boundaries can have large impacts on ecosystem structures and functions at local and regional scales. In this article, we develop a simple model to investigate dynamical implications of bi-directional resource exchanges between two local ecosystems in a meta-ecosystem framework. In our model, we assume that (1) Each local ecosystem acts as both a resource donor and recipient, such that one ecosystem donating resources to another results in a cost to the donating system and a benefit to the recipient; and (2) The costs and benefits of the bi-directional resource exchange between two ecosystems are correlated in a nonlinear fashion. Our model could apply to the resource interactions between terrestrial and aquatic ecosystems that are supported by the literature. Our theoretical results show that bi-directional resource exchange between two ecosystems can indeed generate complicated dynamical outcomes, including the coupled ecosystems having amensalistic, antagonistic, competitive, or mutualistic interactions, with multiple alternative stable states depending on the relative costs and benefits. In addition, if the relative cost for resource exchange for an ecosystem is decreased or the relative benefit for resource exchange for an ecosystem is increased, the production of that ecosystem would increase; however, depending on the local environment, the production of the other ecosystem may increase or decrease. We expect that our work, by evaluating the potential outcomes of resource exchange theoretically, can facilitate empirical evaluations and advance the understanding of spatial ecosystem ecology where resource exchanges occur in varied ecosystems through a complicated network.

The Hyalella (Crustacea: Amphipoda) species cloud of the ancient Lake Titicaca originated from multiple colonizations.

Ancient lakes are renowned for their exceptional diversity of endemic species. As model systems for the study of sympatric speciation, it is necessary to understand whether a given hypothesized species flock is of monophyletic or polyphyletic origin. Here, we present the first molecular characterization of the Hyalella (Crustacea: Amphipoda) species complex of Lake Titicaca, using COI and 28S DNA sequences, including samples from the connected Small and Large Lakes that comprise Lake Titicaca as well as from a broader survey of southern South American sites. At least five evolutionarily distant lineages are present within Lake Titicaca, which were estimated to have diverged from one another 12-20 MYA. These major lineages are dispersed throughout the broader South American Hyalella phylogeny, with each lineage representing at least one independent colonization of the lake. Moreover, complex genetic relationships are revealed between Lake Titicaca individuals and those from surrounding water bodies, which may be explained by repeated dispersal into and out of the lake, combined with parallel intralacustrine diversification within two separate clades. Although further work in deeper waters will be required to determine the number of species present and modes of diversification, our results strongly indicate that this amphipod species cloud is polyphyletic with a complex geographic history.

Synthesis: comparing effects of resource and consumer fluxes into recipient food webs using meta-analysis.

Here we synthesize empirical research using meta-analysis to compare how consumer and resource fluxes affect recipient food webs. We tested the following hypotheses: (H1) The direct effects of resource fluxes (bottom-up) should be stronger than the direct effects of consumer fluxes (top-down), because resource fluxes are permanent (do not return to the food web in which they were produced) but consumer fluxes may not be (consumers can leave). (H2) Following H1, the indirect effects should attenuate (weaken) more quickly for consumer fluxes than for resource fluxes due to their direct effects being weaker, (H3) The effects of resource fluxes should be stronger when recipient food webs are in different ecosystems than donor food webs due to differences in elevation that accompany cross-ecosystem food web interfaces, often increasing flux quantity due to gravity, while the effects of consumer fluxes should be stronger when donor and recipient food webs are in the same ecosystem as they should more easily assimilate into the recipient food web. We found no differences in the magnitude of bottom-up and top-down direct effects for resource and consumer fluxes, but top-down direct effects were 122% stronger than top-down indirect effects. Indirect effects of prey and predator fluxes quickly attenuated while indirect effects of non-prey resource and herbivore fluxes did not, as the overall direct effects of prey and predator fluxes were 123% and 163% stronger than their indirect effects, respectively. This result suggests that the magnitude of indirect effects decrease as the trophic level of resource and consumer fluxes increases, and also contrasts with results from studies showing in situ top-down indirect effects are stronger than in situ bottom-up indirect effects. We found that resource and consumer flux effect sizes were similar when they occurred between ecosystems, but when they occurred within ecosystems predator flux effects were 107% stronger than nutrient flux effects. Finally, we found that observational studies had higher effect sizes than manipulative studies. Future research should focus on how resource and consumer fluxes might interact and generate feedbacks in empirical studies of natural food webs, and what ecological factors might affect their relative strength.

Plant biodiversity effects in reducing fluvial erosion are limited to low species richness.

It has been proposed that plant biodiversity may increase the erosion resistance of soils, yet direct evidence for any such relationship is lacking. We conducted a mesocosm experiment with eight species of riparian herbaceous plants, and found evidence that plant biodiversity significantly reduced fluvial erosion rates, with the eight-species polyculture decreasing erosion by 23% relative to monocultures. Species richness effects were largest at low levels of species richness, with little increase between four and eight species. Our results suggest that plant biodiversity reduced erosion rates indirectly through positive effects on root length and number of root tips, and that interactions between legumes and non-legumes were particularly important in producing biodiversity effects. Presumably, legumes increased root production of non-legumes by increasing soil nitrogen availability due to their ability to fix atmospheric nitrogen. Our data suggest that a restoration project using species from different functional groups might provide the best insurance to maintain long-term erosion resistance.

A tale of two rivers: implications of water management practices for mussel biodiversity outcomes during droughts.

Droughts often pose situations where stream water levels are lowest while human demand for water is highest. Here we present results of an observational study documenting changes in freshwater mussel communities in two southern US rivers during a multi-year drought. During a 13-year period water releases into the Kiamichi River from an impoundment were halted during droughts, while minimum releases from an impoundment were maintained in the Little River. The Kiamichi observed nearly twice as many low-flow events known to cause mussel mortality than the Little, and regression tree analyses suggest that this difference was influenced by reduced releases. During this period mussel communities in the Kiamichi declined in species richness and abundance, changes that were not observed in the Little. These results suggest that reduced releases during droughts likely led to mussel declines in one river, while maintaining reservoir releases may have sustained mussel populations in another.

Bottom-up biodiversity effects increase resource subsidy flux between ecosystems.

Although biodiversity can increase ecosystem productivity and adjacent ecosystems are often linked by resource flows between them, the relationship between biodiversity and resource subsidies is not well understood. Here we test the influence of biodiversity on resource subsidy flux by manipulating freshwater mussel species richness and documenting the effects on a trophic cascade from aquatic to terrestrial ecosystems. In a mesocosm experiment, mussel effects on algae were linked through stable isotope analyses to mussel-derived nitrogen subsidies, but mussel biodiversity effects on algal accumulation were not significant. In contrast, mussel biodiversity significantly increased aquatic insect emergence rates, because aquatic insects were responding to mussel-induced changes in algal community structure instead of algal accumulation. In turn, mussel biodiversity also significantly increased terrestrial spider abundance as spiders tracked increases in aquatic insect prey after a reproduction event. In a comparative field study, we found that sites with greater mussel species richness had higher aquatic insect emergence rates. These results show that, because food webs in adjacent ecosystems are often linked, biodiversity effects in one ecosystem can influence adjacent ecosystems as well.

Density-dependent biodiversity effects on physical habitat modification by freshwater bivalves.

Several decades of research have shown that biodiversity affects ecosystem processes associated with resource capture and the production of biomass within trophic levels. Although there are good reasons to expect that biodiversity influences non-trophic ecosystem processes, such as the physical creation or modification of habitat, studies investigating the role of biodiversity on physical processes are scarce. Here we report the results of a study using artificial streams to test the influence of freshwater mussel biodiversity on gravel erosion during high flows while manipulating mussel abundance. Mussel species vary in traits that should influence their effects on erosion, such as size, shell morphology, and burrowing behavior. We found that mussel species richness was associated with an increase in erosion at both low and high densities. Planned contrasts showed that the erosion observed in species mixtures was purely additive at low density, indicating that erosion in a species polyculture could routinely be predicted by the performance of monocultures. However, at high density certain combinations of species showed nonadditive effects on erosion, suggesting that organism abundance can fundamentally alter biodiversity effects. Although this may have been a result of altered species interactions at high density, our study design cannot confirm this.