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.

Designing flows to enhance ecosystem functioning in heavily altered rivers.

More than a century of dam construction and water development in the western United States has led to extensive ecological alteration of rivers. Growing interest in improving river function is compelling practitioners to consider ecological restoration when managing dams and water extraction. We developed an Ecological Response Model (ERM) for the Cache la Poudre River, northern Colorado, USA, to illuminate effects of current and possible future water management and climate change. We used empirical data and modeled interactions among multiple ecosystem components to capture system-wide insights not possible with the unintegrated models commonly used in environmental assessments. The ERM results showed additional flow regime modification would further alter the structure and function of Poudre River aquatic and riparian ecosystems due to multiple and interacting stressors. Model predictions illustrated that specific peak flow magnitudes in spring and early summer are critical for substrate mobilization, dynamic channel morphology, and overbank flows, with strong subsequent effects on instream and riparian biota that varied seasonally and spatially, allowing exploration of nuanced management scenarios. Instream biological indicators benefitted from higher and more stable base flows and high peak flows, but stable base flows with low peak flows were only half as effective to increase indicators. Improving base flows while reducing peak flows, as currently proposed for the Cache la Poudre River, would further reduce ecosystem function. Modeling showed that even presently depleted annual flow volumes can achieve substantially different ecological outcomes in designed flow scenarios, while still supporting social demands. Model predictions demonstrated that implementing designed flows in a natural pattern, with attention to base and peak flows, may be needed to preserve or improve ecosystem function of the Poudre River. Improved regulatory policies would include preservation of ecosystem-level, flow-related processes and adaptive management when water development projects are considered.

Irrigation canals are newly created streams of semi-arid agricultural regions.

The natural hydrologic processes that create and maintain the diversity of aquatic and riparian habitats along the World's streams and rivers have been profoundly altered by humans. Diversion of surface water to support production agriculture in arid and semi-arid regions has degraded ecosystems but also created potential habitat along and in canals specifically designed to transport water. The prevalence of canals and the immense amount of water used for agriculture have created these new artificial stream systems. This study demonstrates the potential for irrigation canals to support riparian and aquatic communities similar to natural streams in urban/residential and agricultural landscapes. We examined the hydrological and ecological characteristics of streams and irrigation canals in urban and agricultural landscapes in northeastern Colorado, typical of regions dominated by irrigation-supported agriculture. Flow patterns in canals depended on their size and had a range of patterns with potential ecological consequences such as rapidly rising and falling water stage, intermittent dry periods, and delayed peak and base flows compared to natural streams. Despite these hydrologic differences, the taxonomic and functional composition of riparian plant and aquatic macroinvertebrate communities indicated that ecological similarities exist between streams and canals, but are dependent, in part, on their landscape setting with stronger similarities in agricultural areas. We also tested the influence of characterizing taxa by functional groups using physiology, ecology and life history traits to explore attributes of habitats including woody canopy structure and water quality. We used a Habitat Quality Index (HQI) that combined physical and biological measures into a single index. Streams scored higher on average within agriculture and urban/residential settings compared to canals; however, one third of urban canals scored above the average of agricultural streams. This multidisciplinary study shows that irrigation canals can be valuable riparian and aquatic habitat, especially in regions with severely degraded streams.

Flow regime alteration degrades ecological networks in riparian ecosystems.

Riverine ecosystems are governed by patterns of temporal variation in river flows. This dynamism will change due to climate change and the near-ubiquitous human control of river flows globally, which may have severe effects on species distributions and interactions. We employed a combination of population modelling and network theory to explore the consequences of possible flow regime futures on riparian plant communities, including scenarios of increased drought, flooding and flow homogenization (removal of flow variability). We found that even slight modifications to the historic natural flow regime had significant consequences for the structure of riparian plant networks. Networks of emergent interactions between plant guilds were most connected at the natural flow regime and became simplified with increasing flow alteration. The most influential component of flow alteration was flood reduction, with drought and flow homogenization both having greater simplifying community-wide consequences than increased flooding. These findings suggest that maintaining floods under future climates will be needed to overcome the negative long-term consequences of flow modification on riverine ecosystems.

Variation in species-level plant functional traits over wetland indicator status categories.

Wetland indicator status (WIS) describes the habitat affinity of plant species and is used in wetland delineations and resource inventories. Understanding how species-level functional traits vary across WIS categories may improve designations, elucidate mechanisms of adaptation, and explain habitat optima and niche. We investigated differences in species-level traits of riparian flora across WIS categories, extending their application to indicate hydrologic habitat. We measured or compiled data on specific leaf area (SLA), stem specific gravity (SSG), seed mass, and mature height of 110 plant species that occur along the Colorado River in Grand Canyon, Arizona. Additionally, we measured leaf δ13C, δ15N, % carbon, % nitrogen, and C/N ratio of 56 species with C3 photosynthesis. We asked the following: (i) How do species-level traits vary over WIS categories? (ii) Does the pattern differ between herbaceous and woody species? (iii) How well do multivariate traits define WIS categories? (iv) Which traits are correlated? The largest trait differences among WIS categories for herbaceous species occurred for SSG, seed mass, % leaf carbon and height, and for woody species occurred for height, SSG, and δ13C. SSG increased and height decreased with habitat aridity for both woody and herbaceous species. The δ13C and hence water use efficiency of woody species increased with habitat aridity. Water use efficiency of herbaceous species increased with habitat aridity via greater occurrence of C4 grasses. Multivariate trait assemblages differed among WIS categories. Over all species, SLA was correlated with height, δ13C, % leaf N, and C/N; height was correlated with SSG and % leaf C; SSG was correlated with % leaf C. Adaptations of both herbaceous and woody riparian species to wet, frequently inundated habitats include low-density stem tissue. Adaptations to drier habitats in the riparian zone include short, high-density cavitation-resistant stem tissue, and high water use efficiency. The results enhance understanding about using traits to describe plant habitat in riparian systems.

Linking river flow regimes to riparian plant guilds: a community-wide modeling approach.

Modeling riparian plant dynamics along rivers is complicated by the fact that plants have different edaphic and hydrologic requirements at different life stages. With intensifying human demands for water and continued human alteration of rivers, there is a growing need for predicting responses of vegetation to flow alteration, including responses related to climate change and river flow management. We developed a coupled structured population model that combines stage-specific responses of plant guilds with specific attributes of river hydrologic regime. The model uses information on the vital rates of guilds as they relate to different hydrologic conditions (flood, drought, and baseflow), but deliberately omits biotic interactions from the structure (interaction neutral). Our intent was to (1) consolidate key vital rates concerning plant population dynamics and to incorporate these data into a quantitative framework, (2) determine whether complex plant stand dynamics, including biotic interactions, can be predicted from basic vital rates and river hydrology, and (3) project how altered flow regimes might affect riparian communities. We illustrated the approach using five flow-response guilds that encompass much of the river floodplain community: hydroriparian tree, xeroriparian shrub, hydroriparian shrub, mesoriparian meadow, and desert shrub. We also developed novel network-based tools for predicting community-wide effects of climate-driven shifts and deliberately altered flow regimes. The model recovered known patterns of hydroriparian tree vs. xeroriparian shrub dominance, including the relative proportion of these two guilds as a function of river flow modification. By simulating flow alteration scenarios ranging from increased drought to shifts in flood timing, the model predicted that mature hydroriparian forest should be most abundant near the observed natural flow regime. Multiguild sensitivity analysis identified substantial network connectivity (many connected nodes) and biotic linkage (strong pairwise connections between nodes) under natural flow regime conditions. Both connectivity and linkage were substantially reduced under drought and other flow-alteration scenarios, suggesting that community structure is destabilized under such conditions. This structured population modeling approach provides a useful tool for understanding the community-wide effects of altered flow regimes due to climate change and management actions that influence river flow regime.

Linking stream flow and groundwater to avian habitat in a desert riparian system.

Increasing human populations have resulted in aggressive water development in arid regions. This development typically results in altered stream flow regimes, reduced annual flow volumes, changes in fluvial disturbance regimes, changes in groundwater levels, and subsequent shifts in ecological patterns and processes. Balancing human demands for water with environmental requirements to maintain functioning ecosystems requires quantitative linkages between water in streams and ecosystem attributes. Streams in the Sonoran Desert provide important habitat for vertebrate species, including resident and migratory birds. Habitat structure, food, and nest-building materials, which are concentrated in riparian areas, are provided directly or indirectly by vegetation. We measured riparian vegetation, groundwater and surface water, habitat structure, and bird occurrence along Cherry Creek, a perennial tributary of the Salt River in central Arizona, USA. The purpose of this work was to develop an integrated model of groundwater-vegetation-habitat structure and bird occurrence by: (1) characterizing structural and provisioning attributes of riparian vegetation through developing a bird habitat index (BHI), (2) validating the utility of our BHI through relating it to measured bird community composition, (3) determining the riparian plant species that best explain the variability in BHI, (4) developing predictive models that link important riparian species to fluvial disturbance and groundwater availability along an arid-land stream, and (5) simulating the effects of changes in flow regime and groundwater levels and determining their consequences for riparian bird communities. Riparian forest and shrubland vegetation cover types were correctly classified in 83% of observations as a function of fluvial disturbance and depth to water table. Groundwater decline and decreased magnitude of fluvial disturbance caused significant shifts in riparian cover types from riparian forest to shrublands. Variability in the BHI was best explained by the cover of deciduous riparian tree species, primarily Populus fremontii, Platanus wrightii, and Salix gooddingii. The distributions of these plant species were well explained by the depth to groundwater and magnitude of fluvial disturbance along the stream. Bird species diversity and richness were significantly higher in sites with higher habitat indices. This quantitative linkage between surface and groundwater, plant species composition, habitat complexity, and bird communities has implications for water management and in determining environmental flows.

The role of hydrochory in structuring riparian and wetland vegetation.

Hydrochory, or the passive dispersal of organisms by water, is an important means of propagule transport, especially for plants. During recent years, knowledge about hydrochory and its ecological consequences has increased considerably and a substantial body of literature has been produced. Here, we review this literature and define the state of the art of the discipline. A substantial proportion of species growing in or near water have propagules (fruits, seeds or vegetative units) able to disperse by water, either floating, submerged in flowing water, or with the help of floating vessels. Hydrochory can enable plants to colonize sites out of reach with other dispersal vectors, but the timing of dispersal and mechanisms of establishment are important for successful establishment. At the population level, hydrochory may increase the effective size and longevity of populations, and control their spatial configuration. Hydrochory is also an important source of species colonizing recruitment-limited riparian and wetland communities, contributing to maintenance of community species richness. Dispersal by water may even influence community composition in different landscape elements, resulting in landscape-level patterns. Genetically, hydrochory may reduce spatial aggregation of genetically related individuals, lead to high gene flow among populations, and increase genetic diversity in populations receiving many propagules. Humans have impacted hydrochory in many ways. For example, dams affect hydrochory by reducing peak flows and hence dispersal capacity, altering the timing of dispersal, and by presenting physical barriers to dispersal, with consequences for riverine plant communities. Hydrochory has been inferred to be an important vector for the spread of many invasive species, but there is also the potential for enhancing ecosystem restoration by improving or restoring water dispersal pathways. Climate change may alter the role of hydrochory by modifying the hydrology of water-bodies as well as conditions for propagule release and plant colonization.

Shifting dominance of riparian Populus and Tamarix along gradients of flow alteration in western North American rivers.

Tamarix ramosissima is a naturalized, nonnative plant species which has become widespread along riparian corridors throughout the western United States. We test the hypothesis that the distribution and success of Tamarix result from human modification of river-flow regimes. We conducted a natural experiment in eight ecoregions in arid and semiarid portions of the western United States, measuring Tamarix and native Populus recruitment and abundance at 64 sites along 13 perennial rivers spanning a range of altered flow regimes. We quantified biologically relevant attributes of flow alteration as an integrated measure (the index of flow modification, IFM), which was then used to explain between-site variation in abundance and recruitment of native and nonnative riparian plant species. We found the likelihood of successful recruitment of Tamarix to be highest along unregulated river reaches and to remain high across a gradient of regulated flows. Recruitment probability for Populus, in contrast, was highest under free-flowing conditions and declined abruptly under even slight flow modification (IFM > 0.1). Adult Tamarix was most abundant at intermediate levels of IFM. Populus abundance declined sharply with modest flow regulation (IFM > 0.2) and was not present at the most flow-regulated sites. Dominance of Tamarix was highest along rivers with the most altered flow regimes. At the 16 least regulated sites, Tamarix and Populus were equally abundant. Given observed patterns of Tamarix recruitment and abundance, we infer that Tamarix would likely have naturalized, spread, and established widely in riparian communities in the absence of dam construction, diversions, and flow regulation in western North America. However, Tamarix dominance over native species would likely be less extensive in the absence of human alteration of river-flow regimes. Restoration that combines active mechanical removal of established stands of Tamarix with a program of flow releases conducive to native species establishment and persistence is hypothesized to facilitate the codominance of Populus in reaches where it has become rare. Our findings have implications for planning flow-related stream restoration, for developing realistic expectations for yield on investment in prescribed flow releases, and for planning flow-related interventions that might be possible if control and management of invasive plant species along rivers is a goal.

Homogenization of regional river dynamics by dams and global biodiversity implications.

Global biodiversity in river and riparian ecosystems is generated and maintained by geographic variation in stream processes and fluvial disturbance regimes, which largely reflect regional differences in climate and geology. Extensive construction of dams by humans has greatly dampened the seasonal and interannual streamflow variability of rivers, thereby altering natural dynamics in ecologically important flows on continental to global scales. The cumulative effects of modification to regional-scale environmental templates caused by dams is largely unexplored but of critical conservation importance. Here, we use 186 long-term streamflow records on intermediate-sized rivers across the continental United States to show that dams have homogenized the flow regimes on third- through seventh-order rivers in 16 historically distinctive hydrologic regions over the course of the 20th century. This regional homogenization occurs chiefly through modification of the magnitude and timing of ecologically critical high and low flows. For 317 undammed reference rivers, no evidence for homogenization was found, despite documented changes in regional precipitation over this period. With an estimated average density of one dam every 48 km of third- through seventh-order river channel in the United States, dams arguably have a continental scale effect of homogenizing regionally distinct environmental templates, thereby creating conditions that favor the spread of cosmopolitan, nonindigenous species at the expense of locally adapted native biota. Quantitative analyses such as ours provide the basis for conservation and management actions aimed at restoring and maintaining native biodiversity and ecosystem function and resilience for regionally distinct ecosystems at continental to global scales.