Strategic restoration planning for land birds in the Colorado River Delta, Mexico.

Ecological restoration is an essential strategy for mitigating the current biodiversity crisis, yet restoration actions are costly. We used systematic conservation planning principles to design an approach that prioritizes restoration sites for birds and tested it in a riparian forest restoration program in the Colorado River Delta. Restoration goals were to maximize the abundance and diversity of 15 priority birds with a variety of habitat preferences. We built abundance models for priority birds based on the current landscape, and predicted bird distributions and relative abundances under a scenario of complete riparian forest restoration throughout our study area. Then, we used Zonation conservation planning software to rank this restored landscape based on core areas for all priority birds. The locations with the highest ranks represented the highest priorities for restoration and were located throughout the river reach. We optimized how much of the available landscape to restore by simulating restoration of the top 10-90% of ranked sites in 10% intervals. We found that total diversity was maximized when 40% of the landscape was restored, and mean relative abundance was maximized when 80% of the landscape was restored. The results suggest that complete restoration is not optimal for this community of priority birds and restoration of approximately 60% of the landscape would provide a balance between maximum relative abundance and diversity. Subsequent planning efforts will combine our results with an assessment of restoration costs to provide further decision support for the restoration-siting process. Our approach can be applied to any landscape-scale restoration program to improve the return on investment of limited economic resources for restoration.

Provenance, genotype, and flooding influence growth and resource acquisition characteristics in a clonal, riparian shrub.

PREMISE: Riparian plants can exhibit intraspecific phenotypic variability across the landscape related to temperature and flooding gradients. Phenotypes that vary across a climate gradient are often partly genetically determined and may differ in their response to inundation. Changes to inundation patterns across a climate gradient could thus result in site-specific inundation responses. Phenotypic variability is more often studied in riparian trees, yet riparian shrubs are key elements of riparian systems and may differ from trees in phenotypic variability and environmental responses. METHODS: We tested whether individuals of a clonal, riparian shrub, Pluchea sericea, collected from provenances spanning a temperature gradient differed in their phenotypes and responses to inundation and to what degree any differences were related to genotype. Plants were subjected to different inundation depths and a subset genotyped. Variables related to growth and resource acquisition were measured and analyzed using hierarchical, multivariate Bayesian linear regressions. RESULTS: Individuals from different provenances differed in their phenotypes, but not in their response to inundation. Phenotypes were not related to provenance temperature but were partially governed by genotype. Growth was more strongly influenced by inundation, while resource acquisition was more strongly controlled by genotype. CONCLUSIONS: Growth and resource acquisition responses in a clonal, riparian shrub are affected by changes to inundation and plant demographics in unique ways. Shrubs appear to differ from trees in their responses to environmental change. Understanding environmental effects on shrubs separately from those of trees will be a key part of evaluating impacts of environmental change on riparian ecosystems.

Response of riparian vegetation to short- and long-term hydrologic variation.

Increasing demand for river water now conflicts with an increasing desire to maintain riparian ecosystems. Efficiently managing river flows for riparian vegetation requires an understanding of the time scale of flow effects, but this information is limited by the absence of long-term studies of vegetation change in response to flow variation. To investigate the influence of short- and long-term flow variability and dam operation on riparian vegetation, we determined the occurrence of 107 plant species in 133 permanent plots of known inundating discharge along the Gunnison River in Colorado on five different occasions between 1990 and 2013. Individual species moved up and down the gradient of inundating discharge coincident with increases and decreases in mean annual flow, and the correlations between flow and species occurrence were strongest when flows were weighted by time before vegetation sampling with a median half-life of 1.5 years. Some tall, rhizomatous, perennial species, however, responded to flows on a longer time scale. Logistic regression of species occurrence showed a significant relation with inundation duration for 70 out of 107 species. Plot species richness and total vegetative cover decreased in association with desiccation at low inundation durations and with fluvial disturbance at high inundation durations. Within-plot similarity in species occurrence between years decreased strongly with increasing inundation duration. Moderate inundation durations were dominated by tall, rhizomatous, perennial herbs, including invasive Phalaris arundinacea (reed canary grass). Over the 23-year study period, species richness declined, and the proportion of rhizomatous perennials increased, consistent with the hypothesis that decreases in flow peaks and increases in low flows caused by flow regulation have decreased establishment opportunities for disturbance-dependent species. In summary, annual-scale changes in vegetation were strongly influenced by flow variation, and decadal-scale changes were influenced by decreases in fluvial disturbance from upstream flow regulation beginning decades prior to the onset of this study.

Does large dam removal restore downstream riparian vegetation diversity? Testing predictions on the Elwha River, Washington, USA.

Large dams and their removal can profoundly affect riparian ecosystems by altering flow and sediment regimes, hydrochory, and landform dynamics, yet few studies have documented these effects on downstream plant communities. Ecological theory and empirical results suggest that by altering disturbance regimes, reducing hydrochory, and shifting communities to later successional stages, dams reduce downstream plant diversity. Dam removal could reverse these processes, but the release of large volumes of sediment could have unexpected, transient effects. Two large dams were removed on the Elwha River in Washington State, USA, from 2011 to 2014, representing an unprecedented opportunity to study large dam removal effects on riparian plant communities. Our research objectives were to determine: (1) whether the Elwha River dams were associated with lower downstream plant diversity and altered species composition across riparian landforms pre-dam removal, and (2) whether dam removal has begun to restore downstream diversity and composition. To address these objectives, we compared plant species richness and community composition in river segments above, below, and between the two dams. Plant communities were sampled twice before (2005 and 2010) and four times after (2013, 2014, 2016, and 2017) the start of dam removal, with 2013 and 2014 sampled while the upstream dam removal was ongoing. Prior to dam removal, native species richness was 41% lower below dams compared with the upstream segment; 6 years after dam removal began, it increased ~31% between the dams, whereas nonnative species richness and cover were not apparently affected by dams or their removal. Deposition caused by large volumes of released reservoir sediment had mixed effects on native species richness (increased on floodplains, decreased elsewhere) in the lowest river segment. Plant community composition was also different downstream from dams compared with the upstream reference, and has changed in downstream floodplains and bars since dam removal. In the long term, we expect that diversity will continue to increase in downstream river segments. Our results provide evidence that (1) large dams reduce downstream native plant diversity, (2) dam removal may restore it, and (3) given the natural dynamics of riparian vegetation, long-term, multiyear before-and-after monitoring is essential for understanding dam removal effects.

Projected warming disrupts the synchrony of riparian seed release and snowmelt streamflow.

Globally, spring phenology and abiotic processes are shifting earlier with warming. Differences in the magnitudes of these shifts may decouple the timing of plant resource requirements from resource availability. In riparian forests across the northern hemisphere, warming could decouple seed release from snowmelt peak streamflow, thus reducing moisture and safe sites for dominant tree recruitment. We combined field observations with climate, hydrology, and phenology models to simulate future change in synchrony of seed release and snowmelt peaks in the South Platte River Basin, Colorado, for three Salicaceae species that dominate western USA riparian forests. Chilling requirements for overcoming winter endodormancy were strongest in Salix exigua, moderately supported for Populus deltoides, and indiscernible in Salix amygdaloides. Ensemble mean projected warming of 3.5°C shifted snowmelt peaks 10-19 d earlier relative to S. exigua and P. deltoides seed release, because decreased winter chilling combined with increased spring forcing limited change in their phenology. By contrast, warming shifted both snowmelt peaks and S. amygdaloides seed release 21 d earlier, maintaining their synchrony. Decoupling of snowmelt from seed release for Salicaceae with strong chilling requirements is likely to reduce resources critical for recruitment of these foundational riparian forests, although the magnitude of future decoupling remains uncertain.

Conceptualizing Ecological Responses to Dam Removal: If You Remove It, What's to Come?

One of the desired outcomes of dam decommissioning and removal is the recovery of aquatic and riparian ecosystems. To investigate this common objective, we synthesized information from empirical studies and ecological theory into conceptual models that depict key physical and biological links driving ecological responses to removing dams. We define models for three distinct spatial domains: upstream of the former reservoir, within the reservoir, and downstream of the removed dam. Emerging from these models are response trajectories that clarify potential pathways of ecological transitions in each domain. We illustrate that the responses are controlled by multiple causal pathways and feedback loops among physical and biological components of the ecosystem, creating recovery trajectories that are dynamic and nonlinear. In most cases, short-term effects are typically followed by longer-term responses that bring ecosystems to new and frequently predictable ecological condition, which may or may not be similar to what existed prior to impoundment.

Regeneration of Salicaceae riparian forests in the Northern Hemisphere: A new framework and management tool.

Human activities on floodplains have severely disrupted the regeneration of foundation riparian shrub and tree species of the Salicaceae family (Populus and Salix spp.) throughout the Northern Hemisphere. Restoration ecologists initially tackled this problem from a terrestrial perspective that emphasized planting. More recently, floodplain restoration activities have embraced an aquatic perspective, inspired by the expanding practice of managing river flows to improve river health (environmental flows). However, riparian Salicaceae species occupy floodplain and riparian areas, which lie at the interface of both terrestrial and aquatic ecosystems along watercourses. Thus, their regeneration depends on a complex interaction of hydrologic and geomorphic processes that have shaped key life-cycle requirements for seedling establishment. Ultimately, restoration needs to integrate these concepts to succeed. However, while regeneration of Salicaceae is now reasonably well-understood, the literature reporting restoration actions on Salicaceae regeneration is sparse, and a specific theoretical framework is still missing. Here, we have reviewed 105 peer-reviewed published experiences in restoration of Salicaceae forests, including 91 projects in 10 world regions, to construct a decision tree to inform restoration planning through explicit links between the well-studied biophysical requirements of Salicaceae regeneration and 17 specific restoration actions, the most popular being planting (in 55% of the projects), land contouring (30%), removal of competing vegetation (30%), site selection (26%), and irrigation (24%). We also identified research gaps related to Salicaceae forest restoration and discuss alternative, innovative and feasible approaches that incorporate the human component.

Landscape context and the biophysical response of rivers to dam removal in the United States.

Dams have been a fundamental part of the U.S. national agenda over the past two hundred years. Recently, however, dam removal has emerged as a strategy for addressing aging, obsolete infrastructure and more than 1,100 dams have been removed since the 1970s. However, only 130 of these removals had any ecological or geomorphic assessments, and fewer than half of those included before- and after-removal (BAR) studies. In addition, this growing, but limited collection of dam-removal studies is limited to distinct landscape settings. We conducted a meta-analysis to compare the landscape context of existing and removed dams and assessed the biophysical responses to dam removal for 63 BAR studies. The highest concentration of removed dams was in the Northeast and Upper Midwest, and most have been removed from 3rd and 4th order streams, in low-elevation (< 500 m) and low-slope (< 5%) watersheds that have small to moderate upstream watershed areas (10-1000 km2) with a low risk of habitat degradation. Many of the BAR-studied removals also have these characteristics, suggesting that our understanding of responses to dam removals is based on a limited range of landscape settings, which limits predictive capacity in other environmental settings. Biophysical responses to dam removal varied by landscape cluster, indicating that landscape features are likely to affect biophysical responses to dam removal. However, biophysical data were not equally distributed across variables or clusters, making it difficult to determine which landscape features have the strongest effect on dam-removal response. To address the inconsistencies across dam-removal studies, we provide suggestions for prioritizing and standardizing data collection associated with dam removal activities.

Vegetation response to invasive Tamarix control in southwestern U.S. rivers: a collaborative study including 416 sites.

Most studies assessing vegetation response following control of invasive Tamarix trees along southwestern U.S. rivers have been small in scale (e.g., river reach), or at a regional scale but with poor spatial-temporal replication, and most have not included testing the effects of a now widely used biological control. We monitored plant composition following Tamarix control along hydrologic, soil, and climatic gradients in 244 treated and 172 reference sites across six U.S. states. This represents the largest comprehensive assessment to date on the vegetation response to the four most common Tamarix control treatments. Biocontrol by a defoliating beetle (treatment 1) reduced the abundance of Tamarix less than active removal by mechanically using hand and chain-saws (2), heavy machinery (3) or burning (4). Tamarix abundance also decreased with lower temperatures, higher precipitation, and follow-up treatments for Tamarix resprouting. Native cover generally increased over time in active Tamarix removal sites, however, the increases observed were small and was not consistently increased by active revegetation. Overall, native cover was correlated to permanent stream flow, lower grazing pressure, lower soil salinity and temperatures, and higher precipitation. Species diversity also increased where Tamarix was removed. However, Tamarix treatments, especially those generating the highest disturbance (burning and heavy machinery), also often promoted secondary invasions of exotic forbs. The abundance of hydrophytic species was much lower in treated than in reference sites, suggesting that management of southwestern U.S. rivers has focused too much on weed control, overlooking restoration of fluvial processes that provide habitat for hydrophytic and floodplain vegetation. These results can help inform future management of Tamarix-infested rivers to restore hydrogeomorphic processes, increase native biodiversity and reduce abundance of noxious species.

Elevated CO2 does not offset greater water stress predicted under climate change for native and exotic riparian plants.

In semiarid western North American riparian ecosystems, increased drought and lower streamflows under climate change may reduce plant growth and recruitment, and favor drought-tolerant exotic species over mesic native species. We tested whether elevated atmospheric CO2 might ameliorate these effects by improving plant water-use efficiency. We examined the effects of CO2 and water availability on seedlings of two native (Populus deltoides spp. monilifera, Salix exigua) and three exotic (Elaeagnus angustifolia, Tamarix spp., Ulmus pumila) western North American riparian species in a CO2-controlled glasshouse, using 1-m-deep pots with different water-table decline rates. Low water availability reduced seedling biomass by 70-97%, and hindered the native species more than the exotics. Elevated CO2 increased biomass by 15%, with similar effects on natives and exotics. Elevated CO2 increased intrinsic water-use efficiency (Δ¹³C(leaf) ), but did not increase biomass more in drier treatments than wetter treatments. The moderate positive effects of elevated CO2 on riparian seedlings are unlikely to counteract the large negative effects of increased aridity projected under climate change. Our results suggest that increased aridity will reduce riparian seedling growth despite elevated CO2, and will reduce growth more for native Salix and Populus than for drought-tolerant exotic species.

Floristic composition, beta diversity, and nestedness of reference sites for restoration of xeroriparian areas.

In restoration ecology, reference sites serve as models for areas to be restored and can provide a standard of comparison for restoration project outcomes. When reference sites are located a relatively long distance from associated restoration projects, differences in climate, disturbance history, and biogeography can increase beta diversity and may decrease the relevance of reference sites. Variation in factors at the scale of individual reference sites such as patch size, microclimate, barriers to dispersal, or soil chemistry can result in reference site species composition that is a nested subset of the regional species pool. In the Western United States, restoration of riparian areas, particularly those occupied by Tamarix spp., has become a priority; however, little is known about suitable native replacement vegetation communities for relatively dry and saline riparian terraces that comprise many of the sites where Tamarix is removed prior to restoration activities. We studied plant communities on riparian terraces along five rivers in New Mexico, USA, to (1) determine whether the floristic composition of reference sites can be predicted by easily measured soil variables such as pH, salinity (electric conductivity), and texture; (2) examine the extent of distance decay in the compositional similarity of xeroriparian plant communities in the southwestern United States; and (3) determine the degree of nestedness in xeroriparian plant communities in relationship to soil variables. We found that sites clustered into groups based largely on variation in soil salinity and texture. Vegetation across all sites was highly nested with dominant, salt-tolerant species found on most soil groups and salt-intolerant subordinate species restricted to low-salinity soils. The identity of subordinate species was largely site dependent, causing all sites to have the same low degree of similarity regardless of the distance between them. We conclude that, when planning restoration projects on dry and saline riparian sites, soil salinity and texture are good predictors of which species will be most suited to the area being restored, but a candidate species pool should be developed from the nearest possible reference sites, particularly for subordinate species.

Early vegetation development on an exposed reservoir: implications for dam removal.

The 4-year drawdown of Horsetooth Reservoir, Colorado, for dam maintenance, provides a case study analog of vegetation response on sediment that might be exposed from removal of a tall dam. Early vegetation recovery on the exposed reservoir bottom was a combination of (1) vegetation colonization on bare, moist substrates typical of riparian zones and reservoir sediment of shallow dams and (2) a shift in moisture status from mesic to the xeric conditions associated with the pre-impoundment upland position of most of the drawdown zone. Plant communities changed rapidly during the first four years of exposure, but were still substantially different from the background upland plant community. Predictions from the recruitment box model about the locations of Populus deltoides subsp. monilifera (plains cottonwood) seedlings relative to the water surface were qualitatively confirmed with respect to optimum locations. However, the extreme vertical range of water surface elevations produced cottonwood seed regeneration well outside the predicted limits of drawdown rate and height above late summer stage. The establishment and survival of cottonwood at high elevations and the differences between the upland plant community and the community that had developed after four years of exposure suggest that vegetation recovery following tall dam removal will follow a trajectory very different from a simple reversal of the response to dam construction, involving not only long time scales of establishment and growth of upland vegetation, but also possibly decades of persistence of legacy vegetation established during the reservoir to upland transition.

Control of Tamarix in the Western United States: implications for water salvage, wildlife use, and riparian restoration.

Non-native shrub species in the genus Tamarix (saltcedar, tamarisk) have colonized hundreds of thousands of hectares of floodplains, reservoir margins, and other wetlands in western North America. Many resource managers seek to reduce saltcedar abundance and control its spread to increase the flow of water in streams that might otherwise be lost to evapotranspiration, to restore native riparian (streamside) vegetation, and to improve wildlife habitat. However, increased water yield might not always occur and has been substantially lower than expected in water salvage experiments, the potential for successful revegetation is variable, and not all wildlife taxa clearly prefer native plant habitats over saltcedar. As a result, there is considerable debate surrounding saltcedar control efforts. We review the literature on saltcedar control, water use, wildlife use, and riparian restoration to provide resource managers, researchers, and policy-makers with a balanced summary of the state of the science. To best ensure that the desired outcomes of removal programs are met, scientists and resource managers should use existing information and methodologies to carefully select and prioritize sites for removal, apply the most appropriate and cost-effective control methods, and then rigorously monitor control efficacy, revegetation success, water yield changes, and wildlife use.

Germination and Establishment of the Native Plains Cottonwood (Populus deltoides Marshall subsp. monilifera) and the Exotic Russian-olive (Elaeagnus angustifolia L.).

Russian-olive (Elaeagnus angustifolia) is a small Eurasian tree that has escaped from cultivation and become naturalized, primarily along watercourses throughout the western United States. We examined germination and establishment of Russian-olive and plains cottonwood (Populus deltoides), the principal native riparian tree of the Great Plains, under a range of experimental moisture and light conditions. The fewest seedings established under the driest conditions; seedling biomass was predictably lower in the shade; root-to-shoot ratios were higher for cottonwood, higher in the sun, and higher under drier conditions. Several interactions were also significant. The timing of germination and mortality varied between plains cottonwood and Russian-olive: cottonwood germinated in mid-June in all treatments in a single pulse with subsequent mortality; the timing and amount of Russian-olive germination differed substantially across treatments with little net mortality. Differences in life-history traits of these species, including seed size, viability, and dispersal, help explain treatment differences. Russian-olive will likely remain an important component of riparian communities along both unregulated and regulated western rivers because it succeeds under conditions optimal for cottonwood establishment and under many conditions unfavorable for cottonwood. Furthermore, many western states still encourage planting of Russian-olive, and control techniques tend to be labor-intensive and expensive. Germinación y establecimiento del álamo nativo de las planicies (Populus deltoides Marshall subsp. monolifera) y el olivo ruso exótico (Elaeagnus angustifolia L.).

Resumen: El olivo ruso (Elaeagnus angustifolia) es un árbol pequeño nativo de Europa y Asia que ha escapado de ser cultivado y se ha naturalizado principalmente a lo largo de cursos de agua en el oeste de los Estados Unidos. Examinamos la germinación y el establecimiento del olivo ruso y del álamo del llano (Populus deltoides), el principal árbol nativo rivereño de las Grandes Planicies, bajo un amplio espectro de condiciones experimentales de humedad y luz. La menor cantidad de retoños se estableció bajo las condiciones más secas; la biomasa de los retoños fue predeciblemente menor en la sombra; la proporcián raiz; vástago fue más alta para el álamo, mayor bajo el sol y más alta bajo condiciones más secas. También fueron significativas varias interacciones. El tiempo de germinación y la mortalidad para el álamo del llano y el olivo ruso fue diferente. El álamo del llano germinó durante un único pulso a mediados de Junio en todos los tratamientos con la mortalidad subsiguiente, mientras que los tiempos y la cantidad de germinación del olivo ruso difirieron substancialmente entre los distintos tratamientos con muy poca mortalidad neta. Diferencias en los caracteres de la historia de vida de estas especies, incluyendo el tamaño de la semilla, la viabilidad y la dispersión ayuda a explicar las diferencias entre tratamientos. El olivo ruso permanecerá probablemente como un componente importante de las comunidades ribereñas a lo largo de ríos regulados y no-regulados del oeste. Esto es posible porque el olivo ruso se desempeña exitosamente bajo condiciones que son óptimas para el establecimiento del álamo del llano y bajo condiciones diversas desfavorables para álamo. Más el aún, varios estados del oeste todavía fomentan la plantación del olivo ruso y las ténicas de control requieren una labor intensiva y costosa.