Integrated ecosystems: linking food webs through reciprocal resource reliance.

Ecosystems are defined, studied, and managed according to boundaries constructed to conceptualize patterns of interest at a certain scale and scope. The distinction between ecosystems becomes obscured when resources from multiple origins cross porous boundaries and are assimilated into food webs through repeated trophic transfers. Ecosystem compartments can define bounded localities in a heterogeneous landscape that simultaneously retain and exchange energy in the form of organic matter. Here we developed and tested a framework to quantify reciprocal reliance on cross-boundary resource exchange and calculate the contribution of primary production from adjacent ecosystem compartments cycling through food webs to support consumers at different trophic levels. Under this framework, an integrated ecosystem can be measured and designated when the boundary between spatially distinct compartments is permeable and the bidirectional exchange of resources contributes significantly to sustaining both food webs. Using a desert river and riparian zone as a case study, we demonstrate that resources exchanged across the aquatic-riparian boundary cycle through multiple trophic levels. Furthermore, predators on both sides of the boundary were supported by externally produced resources to a similar extent, indicating this is a tightly integrated river-riparian ecosystem and that changes to either compartment will substantially impact the other. Using published data on lake ecosystems, we demonstrated that benthic and pelagic ecosystem compartments are likely not fully integrated, but differences between lakes could be used to test ecological hypotheses. Finally, we discuss how the integrated ecosystem framework could be applied in urban-preserve and field-forest ecosystems to address a broad range of ecological concepts. Because few systems function in complete isolation, this novel approach has application to research and management strategies globally as ecosystems continue to face novel pressures that precipitate cascading ecological repercussions well beyond a bounded system of focus.

Response to Comment on "Designing river flows to improve food security futures in the Lower Mekong Basin".

Williams et al claim that the data used in Sabo et al were improperly scaled to account for fishing effort, thereby invalidating the analysis. Here, we reanalyze the data rescaled per Williams et al and following the methods in Sabo et al Our original conclusions are robust to rescaling, thereby invalidating the assertion that our original analysis is invalid.

Predator water balance alters intraguild predation in a streamside food web.

Previous work suggests that animal water balance can influence trophic interactions, with predators increasing their consumption of water-laden prey to meet water demands. But it is unclear how the need for water interacts with the need for energy to drive trophic interactions under shifting conditions. Using manipulative field experiments, we show that water balance influences the effects of top predators on prey with contrasting ratios of water and energy, altering the frequency of intraguild predation. Water-stressed top predators (large spiders) negatively affect water-laden basal prey (crickets), especially male prey with higher water content, whereas alleviation of water limitation causes top predators to switch to negatively affecting energy-rich midlevel predators (small spiders). Thus, the relative water and energy content of multiple prey, combined with the water demand of the top predator, influences trophic interactions in ways that can alter the strength of intraguild predation. These findings underscore the need for integration of multiresource approaches for understanding implications of global change for food webs.

The U.S. food-energy-water system: A blueprint to fill the mesoscale gap for science and decision-making.

Food, energy, and water (FEW) are interdependent and must be examined as a coupled natural-human system. This perspective essay defines FEW systems and outlines key findings about them as a blueprint for future models to satisfy six key objectives. The first three focus on linking the FEW production and consumption to impacts on Earth cycles in a spatially specific manner in order to diagnose problems and identify potential solutions. The second three focus on describing the evolution of FEW systems to identify risks, thus empowering the FEW actors to better achieve the goals of resilience and sustainability. Four key findings about the FEW systems that guide future model development are (1) that they engage ecological, carbon, water, and nutrient cycles most powerfully among all human systems; (2) that they operate primarily at a mesoscale best captured by counties, districts, and cities; (3) that cities are hubs within the FEW system; and (4) that the FEW system forms a complex network.

Consumption explains intraspecific variation in nutrient recycling stoichiometry in a desert fish.

Consumer-driven nutrient recycling can have substantial effects on primary production and patterns of nutrient limitation in aquatic ecosystems by altering the rates as well as the relative supplies of the key nutrients nitrogen (N) and phosphorus (P). While variation in nutrient recycling stoichiometry has been well-studied among species, the mechanisms that explain intraspecific variation in recycling N:P are not well-understood. We examined the relative importance of potential drivers of variation in nutrient recycling by the fish Gambusia marshi among aquatic habitats in the Cuatro Ciénegas basin of Coahuila, Mexico. There, G. marshi inhabits warm thermal springs with high predation pressure as well as cooler, surface runoff-fed systems with low predation pressure. We hypothesized that variation in food consumption among these habitats would drive intraspecific differences in excretion rates and N:P ratios. Stoichiometric models predicted that temperature alone should not cause substantial variation in excretion N:P, but that further reducing consumption rates should substantially increase excretion N:P. We performed temperature and diet ration manipulation experiments in the laboratory and found strong support for model predictions. We then tested these predictions in the field by measuring nutrient recycling rates and ratios as well as body stoichiometry of fish from nine sites that vary in temperature and predation pressure. Fish from warm, high-predation sites excreted nutrients at a lower N:P ratio than fish from cool, low-predation sites, consistent with the hypothesis that reduced consumption under reduced predation pressure had stronger consequences for P retention and excretion among populations than did variation in body stoichiometry. These results highlight the utility of stoichiometric models for predicting variation in consumer-driven nutrient recycling within a phenotypically variable species.

Detrimental effects of a novel flow regime on the functional trajectory of an aquatic invertebrate metacommunity.

Novel flow regimes resulting from dam operations and overallocation of freshwater resources are an emerging consequence of global change. Yet, anticipating how freshwater biodiversity will respond to surging flow regime alteration requires overcoming two challenges in environmental flow science: shifting from local to riverscape-level understanding of biodiversity dynamics, and from static to time-varying characterizations of the flow regime. Here, we used time-series methods (wavelets and multivariate autoregressive models) to quantify flow-regime alteration and to link time-varying flow regimes to the dynamics of multiple local communities potentially connected by dispersal (i.e., a metacommunity). We studied the Chattahoochee River below Buford dam (Georgia, U.S.A.), and asked how flow regime alteration by a large hydropower dam may control the long-term functional trajectory of the downstream invertebrate metacommunity. We found that seasonal variation in hydropeaking synchronized temporal fluctuations in trait abundance among the flow-altered sites. Three biological trait states describing adaptation to fast flows benefitted from flow management for hydropower, but did not compensate for declines in 16 "loser" traits. Accordingly, metacommunity-wide functional diversity responded negatively to hydropeaking intensity, and stochastic simulations showed that the risk of functional diversity collapse within the next 4 years would decrease by 17% if hydropeaking was ameliorated, or by 9% if it was applied every other season. Finally, an analysis of 97 reference and 23 dam-affected river sites across the U.S. Southeast suggested that flow variation at extraneous, human-relevant scales (12-hr, 24-hr, 1-week) is relatively common in rivers affected by hydropower dams. This study advances the notion that novel flow regimes are widespread, and simplify the functional structure of riverine communities by filtering out taxa with nonadaptive traits and by spatially synchronizing their dynamics. This is relevant in the light of ongoing and future hydrologic alteration due to climate non-stationarity and the new wave of dams planned globally.

Patterns and drivers of fish extirpations in rivers of the American Southwest and Southeast.

Effective conservation of freshwater biodiversity requires spatially explicit investigations of how dams and hydroclimatic alterations among climate regions may interact to drive species to extinction. We investigated how dams and hydroclimatic alterations interact with species ecological and life history traits to influence past extirpation probabilities of native freshwater fishes in the Upper and Lower Colorado River (CR), Alabama-Coosa-Tallapoosa (ACT), and Apalachicola-Chattahoochee-Flint (ACF) basins. Using long-term discharge data for continuously gaged streams and rivers, we quantified streamflow anomalies (i.e., departure "expected" streamflow) at the sub-basin scale over the past half-century. Next, we related extirpation probabilities of native fishes in both regions to streamflow anomalies, river basin characteristics, species traits, and non-native species richness using binomial logistic regression. Sub-basin extirpations in the Southwest (n = 95 Upper CR, n = 130 Lower CR) were highest in lowland mainstem rivers impacted by large dams and in desert springs. Dampened flow seasonality, increased longevity (i.e., delayed reproduction), and decreased fish egg sizes (i.e., lower parental care) were related to elevated fish extirpation probability in the Southwest. Sub-basin extirpations in the Southeast (ACT n = 46, ACF n = 22) were most prevalent in upland rivers, with flow dependency, greater age and length at maturity, isolation by dams, and greater distance upstream. Our results confirm that dams are an overriding driver of native fish species losses, irrespective of basin-wide differences in native or non-native species richness. Dams and hydrologic alterations interact with species traits to influence community disassembly, and very high extirpation risks in the Southeast are due to interactions between high dam density and species restricted ranges. Given global surges in dam building and retrofitting, increased extirpation risks should be expected unless management strategies that balance flow regulation with ecological outcomes are widely implemented.

Interpreting beta-diversity components over time to conserve metacommunities in highly dynamic ecosystems.

The concept of metacommunity (i.e., a set of local communities linked by dispersal) has gained great popularity among community ecologists. However, metacommunity research mostly addresses questions on spatial patterns of biodiversity at the regional scale, whereas conservation planning requires quantifying temporal variation in those metacommunities and the contributions that individual (local) sites make to regional dynamics. We propose that recent advances in diversity-partitioning methods may allow for a better understanding of metacommunity dynamics and the identification of keystone sites. We used time series of the 2 components of beta diversity (richness and replacement) and the contributions of local sites to these components to examine which sites controlled source-sink dynamics in a highly dynamic model system (an intermittent river). The relative importance of the richness and replacement components of beta diversity fluctuated over time, and sample aggregation led to underestimation of beta diversity by up to 35%. Our literature review revealed that research on intermittent rivers would benefit greatly from examination of beta-diversity components over time. Adequately appraising spatiotemporal variability in community composition and identifying sites that are pivotal for maintaining biodiversity at the landscape scale are key needs for conservation prioritization and planning. Thus, our framework may be used to guide conservation actions in highly dynamic ecosystems when time-series data describing biodiversity across sites connected by dispersal are available.

Animal water balance drives top-down effects in a riparian forest-implications for terrestrial trophic cascades.

Despite the clear importance of water balance to the evolution of terrestrial life, much remains unknown about the effects of animal water balance on food webs. Based on recent research suggesting animal water imbalance can increase trophic interaction strengths in cages, we hypothesized that water availability could drive top-down effects in open environments, influencing the occurrence of trophic cascades. We manipulated large spider abundance and water availability in 20 × 20 m open-air plots in a streamside forest in Arizona, USA, and measured changes in cricket and small spider abundance and leaf damage. As expected, large spiders reduced both cricket abundance and herbivory under ambient, dry conditions, but not where free water was added. When water was added (free or within moist leaves), cricket abundance was unaffected by large spiders, but spiders still altered herbivory, suggesting behavioural effects. Moreover, we found threshold-type increases in herbivory at moderately low soil moisture (between 5.5% and 7% by volume), suggesting the possibility that water balance may commonly influence top-down effects. Overall, our results point towards animal water balance as an important driver of direct and indirect species interactions and food web dynamics in terrestrial ecosystems.

Anomalous droughts, not invasion, decrease persistence of native fishes in a desert river.

Changing climate extremes and invasion by non-native species are two of the most prominent threats to native faunas. Predicting the relationships between global change and native faunas requires a quantitative toolkit that effectively links the timing and magnitude of extreme events to variation in species abundances. Here, we examine how discharge anomalies--unexpected floods and droughts--determine covariation in abundance of native and non-native fish species in a highly variable desert river in Arizona. We quantified stochastic variation in discharge using Fourier analyses on >15,000 daily observations. We subsequently coupled maximum annual spectral anomalies with a 15-year time series of fish abundances (1994-2008), using Multivariate Autoregressive State-Space (MARSS) models. Abiotic drivers (discharge anomalies) were paramount in determining long-term fish abundances, whereas biotic drivers (species interactions) played only a secondary role. As predicted, anomalous droughts reduced the abundances of native species, while floods increased them. However, in contrast to previous studies, we observed that the non-native assemblage was surprisingly unresponsive to extreme events. Biological trait analyses showed that functional uniqueness was higher in native than in non-native fishes. We also found that discharge anomalies influenced diversity patterns at the meta-community level, with nestedness increasing after anomalous droughts due to the differential impairment of native species. Overall, our results advance the notion that discharge variation is key in determining community trajectories in the long term, predicting the persistence of native fauna even in the face of invasion. We suggest this variation, rather than biotic interactions, may commonly underlie covariation between native and non-native faunas, especially in highly variable environments. If droughts become increasingly severe due to climate change, and floods increasingly muted due to regulation, fish assemblages in desert rivers may become taxonomically and functionally impoverished and dominated by non-native taxa.

Sensitivity and tolerance of Riparian arthropod communities to altered water resources along a drying river.

BACKGROUND: Rivers around the world are drying with increasing frequency, but little is known about effects on terrestrial animal communities. Previous research along the San Pedro River in southeastern AZ, USA, suggests that changes in the availability of water resources associated with river drying lead to changes in predator abundance, community composition, diversity, and abundance of particular taxa of arthropods, but these observations have not yet been tested manipulatively. METHODS AND RESULTS: In this study, we constructed artificial pools in the stream bed adjacent to a drying section of the San Pedro River and maintained them as the river dried. We compared pitfall trapped arthropods near artificial pools to adjacent control sites where surface waters temporarily dried. Assemblage composition changed differentially at multiple taxonomic levels, resulting in different assemblages at pools than at control sites, with multiple taxa and richness of carabid beetle genera increasing at pools but not at controls that dried. On the other hand, predator biomass, particularly wolf spiders, and diversity of orders and families were consistently higher at control sites that dried. These results suggest an important role for colonization dynamics of pools, as well as the ability of certain taxa, particularly burrowing wolf spiders, to withstand periods of temporary drying. CONCLUSIONS: Overall, we found some agreement between this manipulative study of water resources and a previous analysis of river drying that showed shifts in composition, changes in diversity, and declines in abundance of certain taxa (e.g. carabid beetles). However, colonization dynamics of pools, as well as compensatory strategies of predatory wolf spiders seem to have led to patterns that do not match previous research, with control sites maintaining high diversity, despite drying. Tolerance of river drying by some species may allow persistence of substantial diversity in the face of short-term drying. The long-term effects of drying remain to be investigated.

Crayfish impact desert river ecosystem function and litter-dwelling invertebrate communities through association with novel detrital resources.

Shifts in plant species distributions due to global change are increasing the availability of novel resources in a variety of ecosystems worldwide. In semiarid riparian areas, hydric pioneer tree species are being replaced by drought-tolerant plant species as water availability decreases. Additionally, introduced omnivorous crayfish, which feed upon primary producers, allochthonous detritus, and benthic invertebrates, can impact communities at multiple levels through both direct and indirect effects mediated by drought-tolerant plants. We tested the impact of both virile crayfish (Orconectes virilis) and litter type on benthic invertebrates and the effect of crayfish on detrital resources across a gradient of riparian vegetation drought-tolerance using field cages with leaf litter bags in the San Pedro River in Southeastern Arizona. Virile crayfish increased breakdown rate of novel drought-tolerant saltcedar (Tamarix ramosissima), but did not impact breakdown of drought-tolerant seepwillow (Baccharis salicifolia) or hydric Fremont cottonwood (Populus fremontii) and Gooding's willow (Salix goodingii). Effects on invertebrate diversity were observed at the litter bag scale, but no effects were found at the cage scale. Crayfish decreased alpha diversity of colonizing macroinvertebrates, but did not affect beta diversity. In contrast, the drought-tolerant litter treatment decreased beta diversity relative to hydric litter. As drought-tolerant species become more abundant in riparian zones, their litter will become a larger component of the organic matter budget of desert streams which may serve to homogenize the litter-dwelling community and support elevated populations of virile crayfish. Through impacts at multiple trophic levels, crayfish have a significant effect on desert stream ecosystems.

Dams in the Cadillac Desert: downstream effects in a geomorphic context.

This paper was motivated by the 25th anniversary of the publication of Marc Reisner's book, Cadillac Desert: The American West and its Disappearing Water. Dams are ubiquitous on rivers in the United States, and large dams and storage reservoirs are the hallmark of western U.S. riverscapes. The effects of dams on downstream river ecosystems have attracted much attention and are encapsulated in the serial discontinuity concept (SDC). In the SDC, dams create abrupt shifts in continua of downstream changes in physical and biotic properties. In this paper, we develop a framework for understanding how channel geometry and network structure influence how the physical components of habitat and the biota rebound from discontinuities set up by large dams. We apply this framework to data describing the flow regime, temperature, sediment flux, and fish community composition below Garrison Dam on the Missouri River, Glen Canyon Dam on the Colorado River, and Flaming Gorge Dam on the Green River. Sediment flux in dam tailwaters is under strong control by channel geometry. By contrast, dam-related changes in temperature and flow variation are not significantly modulated by channel geometry or tributary inputs if flow volumes are small (Missouri and Colorado River tributaries). Instead, small tributaries provide near-native conditions (flow and temperature variation) and, as such, provide key refuges for biota from novel habitats in mainstem rivers below large dams. Unregulated tributaries that are large relative to their respective mainstem (e.g., Yampa River) provide refuges as well as significant amelioration of flow and temperature effects from upstream dams. Finally, the proportion of native fish increases with distance from dam and exhibits sharp increases near tributary junctions. These results suggest that tributaries-even minor ones in terms of relative discharge-act as key refugia for native species in regulated river networks. Moreover, large, unregulated tributaries are key to restoring continuity in physical habitat and the biota in large regulated rivers.

Multitaxonomic diversity patterns along a desert riparian-upland gradient.

Riparian areas are noted for their high biodiversity, but this has rarely been tested across a wide range of taxonomic groups. We set out to describe species richness, species abundance, and community similarity patterns for 11 taxonomic groups (forbs & grasses, shrubs, trees, solpugids, spiders, scarab beetles, butterflies, lizards, birds, rodents, and mammalian carnivores) individually and for all groups combined along a riparian-upland gradient in semiarid southeastern Arizona, USA. Additionally, we assessed whether biological characteristics could explain variation in diversity along the gradient using five traits (trophic level, body size, life span, thermoregulatory mechanism, and taxonomic affiliation). At the level of individual groups diversity patterns varied along the gradient, with some having greater richness and/or abundance in riparian zones whereas others were more diverse and/or abundant in upland zones. Across all taxa combined, riparian zones contained significantly more species than the uplands. Community similarity between riparian and upland zones was low, and beta diversity was significantly greater than expected for most taxonomic groups, though biological traits explained little variance in diversity along the gradient. These results indicate heterogeneity amongst taxa in how they respond to the factors that structure ecological communities in riparian landscapes. Nevertheless, across taxonomic groups the overall pattern is one of greater species richness and abundance in riparian zones, coupled with a distinct suite of species.

A landscape perspective on bat foraging ecology along rivers: does channel confinement and insect availability influence the response of bats to aquatic resources in riverine landscapes?

River and riparian areas provide an important foraging habitat for insectivorous bats owing to high insect availability along waterways. However, structural characteristics of the riverine landscape may also influence the location of foraging bats. We used bat detectors to compare bat activity longitudinally along river reaches with contrasting channel confinement, ratio of valley floor width to active channel width, and riparian vegetation, and laterally with distance from the river along three different reach types. We measured rates of insect emergence from the river and aerial insect availability above the river and laterally up to 50-m into the riparian habitat in order to assess the relationship between food resources and insectivorous bat activity. Longitudinally, bat activity was concentrated along confined reaches in comparison to unconfined reaches but was not related to insect availability. Laterally, bats tracked exponential declines in aquatic insects with distance from the river. These data suggest that along the lateral dimension bats track food resources, but that along the longitudinal dimension channel shape and landscape structure determine bat distributions more than food resources.

Reclaiming freshwater sustainability in the Cadillac Desert.

Increasing human appropriation of freshwater resources presents a tangible limit to the sustainability of cities, agriculture, and ecosystems in the western United States. Marc Reisner tackles this theme in his 1986 classic Cadillac Desert: The American West and Its Disappearing Water. Reisner's analysis paints a portrait of region-wide hydrologic dysfunction in the western United States, suggesting that the storage capacity of reservoirs will be impaired by sediment infilling, croplands will be rendered infertile by salt, and water scarcity will pit growing desert cities against agribusiness in the face of dwindling water resources. Here we evaluate these claims using the best available data and scientific tools. Our analysis provides strong scientific support for many of Reisner's claims, except the notion that reservoir storage is imminently threatened by sediment. More broadly, we estimate that the equivalent of nearly 76% of streamflow in the Cadillac Desert region is currently appropriated by humans, and this figure could rise to nearly 86% under a doubling of the region's population. Thus, Reisner's incisive journalism led him to the same conclusions as those rendered by copious data, modern scientific tools, and the application of a more genuine scientific method. We close with a prospectus for reclaiming freshwater sustainability in the Cadillac Desert, including a suite of recommendations for reducing region-wide human appropriation of streamflow to a target level of 60%.

The role of discharge variation in scaling of drainage area and food chain length in rivers.

Food chain length (FCL) is a fundamental component of food web structure. Studies in a variety of ecosystems suggest that FCL is determined by energy supply, environmental stability, and/or ecosystem size, but the nature of the relationship between environmental stability and FCL, and the mechanism linking ecosystem size to FCL, remain unclear. Here we show that FCL increases with drainage area and decreases with hydrologic variability and intermittency across 36 North American rivers. Our analysis further suggests that hydrologic variability is the mechanism underlying the correlation between ecosystem size and FCL in rivers. Ecosystem size lengthens river food chains by integrating and attenuating discharge variation through stream networks, thereby enhancing environmental stability in larger river systems.

The cost of male aggression and polygyny in California sea lions (Zalophus californianus).

In polygynous mating systems, males often increase their fecundity via aggressive defense of mates and/or resources necessary for successful mating. Here we show that both male and female reproductive behavior during the breeding season (June-August) affect female fecundity, a vital rate that is an important determinant of population growth rate and viability. By using 4 years of data on behavior and demography of California sea lions (Zalophus californianus), we found that male behavior and spatial dynamics--aggression and territory size--are significantly related to female fecundity. Higher rates of male aggression and larger territory sizes were associated with lower estimates of female fecundity within the same year. Female aggression was significantly and positively related to fecundity both within the same year as the behavior was measured and in the following year. These results indicate that while male aggression and defense of territories may increase male fecundity, such interactions may cause a reduction in the overall population growth rate by lowering female fecundity. Females may attempt to offset male-related reductions in female fecundity by increasing their own aggression-perhaps to defend pups from incidental injury or mortality. Thus in polygynous mating systems, male aggression may increase male fitness at the cost of female fitness and overall population viability.

Reduced wind speed improves plant growth in a desert city.

BACKGROUND: The often dramatic effects of urbanization on community and ecosystem properties, such as primary productivity, abundances, and diversity are now well-established. In most cities local primary productivity increases and this extra energy flows upwards to alter diversity and relative abundances in higher trophic levels. The abiotic mechanisms thought to be responsible for increases in urban productivity are altered temperatures and light regimes, and increased nutrient and water inputs. However, another abiotic factor, wind speed, is also influenced by urbanization and well known for altering primary productivity in agricultural systems. Wind effects on primary productivity have heretofore not been studied in the context of urbanization. METHODOLOGY/PRINCIPAL FINDINGS: We designed a field experiment to test if increased plant growth often observed in cities is explained by the sheltering effects of built structures. Wind speed was reduced by protecting Encelia farinosa (brittlebush) plants in urban, desert remnant and outlying desert localities via windbreaks while controlling for water availability and nutrient content. In all three habitats, we compared E. farinosa growth when protected by experimental windbreaks and in the open. E. farinosa plants protected against ambient wind in the desert and remnant areas grew faster in terms of biomass and height than exposed plants. As predicted, sheltered plants did not differ from unprotected plants in urban areas where wind speed is already reduced. CONCLUSION/SIGNIFICANCE: Our results indicate that reductions in wind speed due to built structures in cities contribute to increased plant productivity and thus also to changes in abundances and diversity of higher trophic levels. Our study emphasizes the need to incorporate wind speed in future urban ecological studies, as well as in planning for green space and sustainable cities.