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Bright spots for inland fish and fisheries to guide future hydropower development

Hydropower production is one of the greatest threats to fluvial ecosystems and freshwater biodiversity. Now that we have entered the Anthropocene , there is an opportunity to reflect on what might constitute a ‘sustainable’ Anthropocene in the context of hydropower and riverine fish populations. Considering elements of existing practices that promote favorable social-ecological outcomes (i.e., ‘bright spots’) is timely given that there are plans to expand hydropower capacity in previously undammed rivers, intensify dam development in some of the world's largest river systems, and re-license existing facilities. We approach this from a pragmatic perspective: for the foreseeable future, hydropower will likely remain an important source of renewable electricity. To offer support for moving toward a more ‘sustainable’ Anthropocene, we provide syntheses of best practices during the siting, design, construction, operation, and compensation phases of hydropower development to minimize impacts on inland fish. For each phase, we offer positive examples (or what might be considered ‘bright spots’) pertaining to some of the approaches described within our syntheses, acknowledging that these projects may not be viewed as without ecological and (or) societal detriment by all stakeholders. Our findings underscore the importance of protecting critical habitat and free-flowing river reaches through careful site selection and basin-scale planning, infrastructure designs that minimize reservoir effects and facilitate safe passage of fish, construction of hydropower plants using best practices that minimize long-term damage, operating guidelines that mimic natural flow conditions, and compensation that is lasting, effective, inclusive, and locally relevant. Learning from these ‘bright spots’ may require engagement of diverse stakeholders, professionals, and governments at scales that extend well beyond a given site, river, or even basin. Indeed, environmental planning that integrates hydropower development into broader discussions of conserving regional biodiversity and ecosystem services will be of utmost importance.

Water Biology and Security↗

Environmental conditions influencing the abundance of the salmonid ectoparasite Salmincola californiensis across upper Willamette River Reservoirs, Oregon

The freshwater copepod Salmincola californiensis is an ectoparasite of Pacific salmon and trout ( Oncorhynchus spp.). High levels of infection by this parasite can significantly damage gills and result in blood loss, affecting the fitness and survival of hosts, and it may hinder recovery efforts of threatened and endangered salmonids. Juvenile salmonids that rear in reservoirs have been reported to experience higher rates of infection than their stream-dwelling counterparts. To date, the relationship between reservoir environmental conditions and infection rates remains poorly understood. Here, we evaluated sampling methods designed to capture S. californiensis copepodids, the free-swimming infectious life stage of this parasitic copepod, and develop predictive models of parasite abundance in reservoirs . We used light traps to collect 675 zooplankton samples from various sites and depths in Cougar, Lookout Point, and Fall Creek reservoirs, Oregon, USA during five months in 2020. We tested several predictive models of parasite abundance using environmental covariates obtained concurrently during plankton surveys ( e.g. , temperature, ambient light and water clarity, reservoir plankton profiles, and water flow). Our models showed a strong influence of water temperature on abundance of copepodids, with maximum initial abundance occurring at the mean water temperature 15.2 °C. At that temperature, August abundance was estimated to be 58.6 copepodids per site in Cougar Reservoir, 43.8 copepodids per site in Fall Creek Reservoir, and 3.5 copepodids per site in Lookout Point Reservoir. Water temperature and reservoir outflow both affected population growth of copepodids: increased water temperature was associated with an increase in population growth rate while an increased reservoir outflow was associated with a decrease in population growth rate. The efficacy of our sampling method was influenced by percent of moon fullness, whether the sample site was above or below the thermocline, water temperature, and number of juvenile sculpin fishes ( Cottus spp.) captured in the trap. The mean probability of detecting a single copepodid if one was present was 0.042, but detection probability increased to 0.29 under ideal trap set conditions of warmer water, location above the thermocline, and a full moon. Biologists and managers could use these methods to assess the presence and abundance of S. californiensis in other locations, or to inform reservoir operations to reduce potential salmonid infections.

Oregon↗

Southern Salish Sea Habitat Map Series: Admiralty Inlet

In 2010 the Environmental Protection Agency, Region 10 initiated the Puget Sound Scientific Studies and Technical Investigations Assistance Program, designed to support research in support of implementing the Puget Sound Action Agenda. The Action Agenda was created in response to Puget Sound having been designated as one of 28 estuaries of national significance under section 320 of the U.S. Clean Water Act, and its overall goal is to restore the Puget Sound Estuary's environment by 2020. The Southern Salish Sea Mapping Project was funded by the Assistance Program request for proposals process, which also supports a large number of coastal-zone- and ocean-management issues. The issues include the recommendations of the Marine Protected Areas Work Group to the Washington State Legislature (Van Cleve and others, 2009), which endorses a Puget Sound and coast-wide marine conservation needs assessment, gap analysis of existing Marine Protected Areas (MPA) and recommendations for action. This publication is the first of four U.S. Geological Survey Scientific Investigation Maps that make up the Southern Salish Sea Mapping Project. The remaining three map blocks to be published in the future, located south of Admiralty Inlet, are shown in figure 1. Puget Sound is a deep, fjord-type estuary covering an area of 2,330 km 2 in the Pacific Northwest region of the United States (fig. 1). It is connected to the ocean by the Strait of Juan de Fuca, a turbulent passage approximately 160 km in length and 22 km wide at its west end, expanding to over 40 km wide at its east end (Thomson, 1994). During the Pleistocene, the area was occupied several times by lobes of continental ice, resulting in a complex basin-fill of glacial and interglacial deposits that are locally as thick as 1100 m (Johnson and others, 2001). The last glaciation, called the Fraser glaciation, began after 28,800±740 14 C yr B.P. when ice started a slow expansion (Clague, 1981). At peak advance the westward Juan de Fuca lobe reached the edge of the continental shelf through the Juan de Fuca Strait shortly before 14,460±200 14 C yr B.P. (Herzer and Bornhold, 1982). The southward Puget lobe advanced to its terminal position in Puget Sound by around 14,150 14 C yr B.P. (Porter and Swanson, 1998). Ice retreated from its maximum to northern Whidbey Island by 13,650±350 14 C yr B.P. (Dethier and others, 1995). Retreating glaciers resulted in a thick sequence of ice-contact, glacial-marine sediment, and early post-glacial sediments (Linden and Schurrer, 1988). These deposits have experienced the effects of a marine transgression followed by regression, resulting in a sea-level several tens of meters lower than the present day (Linden and Schurrer, 1988). A second transgression brought sea level to about the present level by around 5,470±120 14 C yr B.P. (Clague and others, 1982) establishing the present oceanographic and geologic environment Puget Sound is separated into four interconnected basins; Whidbey, Central (Main), Hood Canal, and South (Thomson, 1994). The Whidbey, Central, and Hood Canal basins are the three main branches of the Puget Sound estuary and are separated from the Strait of Juan de Fuca by a double sill at Admiralty Inlet. The Admiralty Inlet map area includes the Inlet and a portion of the Whidbey Basin (fig. 1). The shallower South Basin is separated by a sill at Tacoma Narrows and is highly branched with numerous finger inlets. Flow within Puget Sound is dominated by tidal currents of as much as 1 m/s at Admiralty Inlet, reducing to approximately 0.5 m/s in the Central Basin (Lavelle and others, 1988). The lack of silt and clay-sized sediments in the Admiralty Inlet map area is likely a result of the strong currents (see Ground-Truth Studies for the Admiralty Inlet Map Area, sheet 3). The subtidal component of flow reaches approximately 0.1 m/s and is driven by density gradients arising from the contrast in salty ocean water at the entrance and freshwater inputs from stream flow (Lavelle and others, 1988). The total freshwater input to Puget Sound is approximately 3.4 x 10 6 m 3 /day, primarily from the Skagit River (Cannon, 1983). The subtidal circulation mostly consists of a two-layered flow in the basins with fresher water exiting at the surface and saltier water entering at depth (Ebbesmeyer and Cannon, 2001). In general, surface waters flow north and deeper waters flow south; variations arise from wind effects that can drive a surface current in the same direction as the wind, and a baroclinic response in the lower layer to about 100-m depth (Matsuura and Cannon, 1997). Oceanographic properties are influenced by temporal forcing parameters such as reduced stream flow during the 2000-01 drought that increased surface salinity and decreased differences between surface and bottom waters (Newton and others, 2003). On offshore seismic-reflection profiles, Pleistocene strata (excluding latest Pleistocene glacial and post-glacial deposits) form a distinct seismic unit, bounded below by pre-Tertiary or Tertiary basement and above by typically flat-lying latest Pleistocene to Holocene deposits that fill in erosional or depositional relief (Johnson and others, 2001). Cores from central Puget Sound have accumulation rates that range from 85 to 1200 mg/cm 2 /yr, or 0.12 to 2.4 cm/yr; the highest accumulation rates are near the southern end of central Puget Sound (Carpenter and others, 1985). Carpenter and others (1985) un-weighted arithmetic mean of accumulation rates for central Puget Sound deeper stations is 480±340 (± one standard deviation) mg/cm 2 /yr. Lavelle and others (1985) also found rates as high as 1200 mg/cm 2 /yr over the past approximately 70 years in cores in the Central Basin off of and north and south of Elliott Bay. Puget Sound basin rates are comparable to rates in midshelf silt deposits on the Washington coast north of the Columbia River (Nittrouer and others, 1979). The deep subtidal (in other words, below SCUBA depths) habitats of Puget Sound are relatively poorly known. A few subtidal surveys exist for several habitat types from the 1960s and 1970s (reviewed in Dethier, 1990), using grab and box core data. The Dethier (1990) review divides habitat up into Coast and Marine Ecological Classification Standard (CMECS) substrate, water column energy, and depth zones but does not attempt to map these habitats, rather it is an inventory of habitats found in the area and the flora and fauna associated with each habitat. The approach of the Southern Salish Sea Mapping project is to create highly detailed seafloor maps through collection, integration, interpretation, and visualization of swath sonar data (the undersea equivalent of satellite remote-sensing data in terrestrial mapping), acoustic backscatter, seafloor video, seafloor photography, and bottom-sediment sampling data. This approach is based in part on methods presented and data collection and product needs identified at the Washington State Seafloor Mapping Workshop (Washington State Seafloor Mapping Workshop Steering Committee, 2008), attended by coastal and marine managers and scientists. The map products display seafloor geomorphology and substrate, and identify potential marine benthic habitats. It is emphasized that the more interpretive habitat and geology maps rely on the integration of multiple, new high-resolution datasets and that mapping at small scales would not be possible without such data. Oceanographic current and wave data is not included in this analysis, however, the accompanying geographic information system (GIS) data set is designed and intended to be combined with oceanographic and biologic data sets assembled by others in the future and some of the GIS data has already been incorporated in the unpublished Nature Conservancy Benthic Habitats of Puget Sound database. This publication includes four map sheets, explanatory text, and a descriptive pamphlet. Each map sheet is published as a portable document format (PDF) file. ESRI ArcGIS compatible geotiffs (for example, bathymetry) and shapefiles (for example video observation points) will be available for download in the data catalog associated with this publication (Cochrane, 2015). An ArcGIS Project File with the symbology used to generate the map sheets is also provided. For those who do not own the full suite of ESRI GIS and mapping software, the data can be read using ESRI ArcReader, a free viewer that is available at http://www.esri.com/software/arcgis/arcreader/index.html .

Washington↗

Identifying dominant environmental predictors of freshwater wetland methane fluxes across diurnal to seasonal time scales

While wetlands are the largest natural source of methane (CH 4 ) to the atmosphere, they represent a large source of uncertainty in the global CH 4 budget due to the complex biogeochemical controls on CH 4 dynamics. Here we present, to our knowledge, the first multi-site synthesis of how predictors of CH 4 fluxes (FCH4) in freshwater wetlands vary across wetland types at diel, multiday (synoptic), and seasonal time scales. We used several statistical approaches (correlation analysis, generalized additive modeling, mutual information, and random forests) in a wavelet-based multi-resolution framework to assess the importance of environmental predictors, nonlinearities and lags on FCH4 across 23 eddy covariance sites. Seasonally, soil and air temperature were dominant predictors of FCH4 at sites with smaller seasonal variation in water table depth (WTD). In contrast, WTD was the dominant predictor for wetlands with smaller variations in temperature (e.g., seasonal tropical/subtropical wetlands). Changes in seasonal FCH4 lagged fluctuations in WTD by ~17 ± 11 days, and lagged air and soil temperature by median values of 8 ± 16 and 5 ± 15 days, respectively. Temperature and WTD were also dominant predictors at the multiday scale. Atmospheric pressure (PA) was another important multiday scale predictor for peat-dominated sites, with drops in PA coinciding with synchronous releases of CH 4 . At the diel scale, synchronous relationships with latent heat flux and vapor pressure deficit suggest that physical processes controlling evaporation and boundary layer mixing exert similar controls on CH 4 volatilization, and suggest the influence of pressurized ventilation in aerenchymatous vegetation. In addition, 1- to 4-h lagged relationships with ecosystem photosynthesis indicate recent carbon substrates, such as root exudates, may also control FCH4. By addressing issues of scale, asynchrony, and nonlinearity, this work improves understanding of the predictors and timing of wetland FCH4 that can inform future studies and models, and help constrain wetland CH 4 emissions.

Global Change Biology↗

Spatially explicit habitat models for 28 fishes from the Upper Mississippi River System (AHAG 2.0)

Environmental management actions in the Upper Mississippi River System (UMRS) typically require pre-project assessments of predicted benefits under a range of project scenarios. The U.S. Army Corps of Engineers (USACE) now requires certified and peer-reviewed models to conduct these assessments. Previously, habitat benefits were estimated for fish communities in the UMRS using the Aquatic Habitat Appraisal Guide (AHAG v.1.0; AHAG from hereon). This spreadsheet-based model used a habitat suitability index (HSI) approach that drew heavily upon Habitat Evaluation Procedures (HEP; U.S. Fish and Wildlife Service, 1980) by the U.S. Fish and Wildlife Service (USFWS). The HSI approach requires developing species response curves for different environmental variables that seek to broadly represent habitat. The AHAG model uses species-specific response curves assembled from literature values, data from other ecosystems, or best professional judgment. A recent scientific review of the AHAG indicated that the model’s effectiveness is reduced by its dated approach to large river ecosystems, uncertainty regarding its data inputs and rationale for habitat-species response relationships, and lack of field validation (Abt Associates Inc., 2011). The reviewers made two major recommendations: (1) incorporate empirical data from the UMRS into defining the empirical response curves, and (2) conduct post-project biological evaluations to test pre-project benefits estimated by AHAG. Our objective was to address the first recommendation and generate updated response curves for AHAG using data from the Upper Mississippi River Restoration-Environmental Management Program (UMRR-EMP) Long Term Resource Monitoring Program (LTRMP) element. Fish community data have been collected by LTRMP (Gutreuter and others, 1995; Ratcliff and others, in press) for 20 years from 6 study reaches representing 1,930 kilometers of river and >140 species of fish. We modeled a subset of these data (28 different species; occurrences at sampling sites as observed in day electrofishing samples) using multiple logistic regression with presence/absence responses. Each species’ probability of occurrence, at each sample site, was modeled as a function of 17 environmental variables observed at each sample site by LTRMP standardized protocols. The modeling methods used (1) a forward-selection process to identify the most important predictors and their relative contributions to predictions; (2) partial methods on the predictor set to control variance inflation; and (3) diagnostics for LTRMP design elements that may influence model fits. Models were fit for 28 species, representing 3 habitat guilds (Lentic, Lotic, and Generalist). We intended to develop “systemic models” using data from all six LTRMP study reaches simultaneously; however, this proved impossible. Thus, we “regionalized” the models, creating two models for each species: “Upper Reach” models, using data from Pools 4, 8, and 13; and “Lower Reach” models, using data from Pool 26, the Open River Reach of the Mississippi River, and the La Grange reach of the Illinois River. A total of 56 models were attempted. For any given site-scale prediction, each model used data from the three LTRMP study reaches comprising the regional model to make predictions. For example, a site-scale prediction in Pool 8 was made using data from Pools 4, 8, and 13. This is the fundamental nature and trade-off of regionalizing these models for broad management application. Model fits were deemed “certifiably good” using the Hosmer and Lemeshow Goodness-of-Fit statistic (Hosmer and Lemeshow, 2000). This test post-partitions model predictions into 10 groups and conducts inferential tests on correspondences between observed and expected probability of occurrence across all partitions, under Chi-square distributional assumptions. This permits an inferential test of how well the models fit and a tool for reporting when they did not (and perhaps why). Our goal was to develop regionalized models, and to assess and describe circumstances when a good fit was not possible. Seven fish species composed the Lentic guild. Good fits were achieved for six Upper Reach models. In the Lower Reach, no model produced good fits for the Lentic guild. This was due to (1) lentic species being much less prominent in the Lower Reach study areas, and (2) those that do express greater prominence principally do so only in the La Grange reach of the Illinois River. Thus, developing Lower Reach models for Lentic species will require parsing La Grange from the other two Lower Reach study areas and fitting separate models. We did not do that as part of this study, but it could be done at a later time. Nine species comprised the Lotic guild. Good fits were achieved for seven Upper Reach models and six Lower Reach models. Four species had good fits for both regions (flathead catfish, blue sucker, sauger, and shorthead redhorse). Three species showed zoogeographic zonation, with a good model fit in one of the regions, but not in the region in which they were absent or rarely occurred (blue catfish, rock bass, and skipjack herring). Twelve species comprised the Generalist guild. Good fits were achieved for five Upper Reach models and eight Lower Reach models. Six species had good fits for both regions (brook silverside, emerald shiner, freshwater drum, logperch, longnose gar, and white bass). Two species showed zoogeographic zonation, with a good model fit in one of the regions, but not in the region in which they were absent or rarely occurred (red shiner and blackstripe topminnow). Poorly fit models were almost always due to the diagnostic variable “field station,” a surrogate for river mile. In these circumstances, the residuals for “field station” were non-randomly distributed and often strongly ordered. This indicates either fitting “pool scale” models for these species and regions, or explicitly model covariances between “field station” and the other predictors within the existing modeling framework. Further efforts on these models should seek to resolve these issues using one of these two approaches. In total, nine species, representing two of the three guilds (Lotic and Generalist), produced well-fit models for both regions. These nine species should comprise the basis for AHAG 2.0. Additional work, likely requiring downscaling of the regional models to pool-scale models, will be needed to incorporate additional species. Alternately, a regionalized AHAG could be comprised of those species, per region, that achieved well-fit models. The number of species and the composition of the regional species pools will differ among regions as a consequence. Each of these alternatives has both pros and cons, and managers are encouraged to consider them fully before further advancing this approach to modeling multi-species habitat suitability.

Upper Mississippi River System↗

Global conservation status of turtles and tortoises (order Testudines)

We present a review and analysis of the conservation status and International Union for Conservation of Nature (IUCN) threat categories of all 360 currently recognized species of extant and recently extinct turtles and tortoises (Order Testudines). Our analysis is based on the 2018 IUCN Red List status of 251 listed species, augmented by provisional Red List assessments by the IUCN Tortoise and Freshwater Turtle Specialist Group (TFTSG) of 109 currently unlisted species of tortoises and freshwater turtles, as well as re-assessments of several outdated IUCN Red List assessments. Of all recognized species of turtles and tortoises, this combined analysis indicates that 20.0% are Critically Endangered (CR), 35.3% are Critically Endangered or Endangered (CR+EN), and 51.9% are Threatened (CR+EN+Vulnerable). Adjusting for the potential threat levels of Data Deficient (DD) species indicates that 56.3% of all data-sufficient species are Threatened. We calculated percentages of imperiled species and modified Average Threat Levels (ATL; ranging from Least Concern = 1 to Extinct = 8) for various taxonomic and geographic groupings. Proportionally more species in the subfamily Geoemydinae (Asian members of the family Geoemydidae) are imperiled (74.2% CR+EN, 79.0% Threatened, 3.89 ATL) compared to other taxonomic groupings, but the families Podocnemididae, Testudinidae, and Trionychidae and the superfamily Chelonioidea (marine turtles of the families Cheloniidae and Dermochelyidae) also have high percentages of imperiled species and ATLs (42.9–50.0% CR+EN, 73.8–100.0% Threatened, 3.44–4.06 ATL). The subfamily Rhinoclemmydinae (Neotropical turtles of the family Geoemydidae) and the families Kinosternidae and Pelomedusidae have the lowest percentages of imperiled species and ATLs (0%–7.4% CR+EN, 7.4%–13.3% Threatened, 1.65–1.87 ATL). Turtles from Asia have the highest percentages of imperiled species (75.0% CR+EN, 83.0% Threatened, 3.98 ATL), significantly higher than predicted based on the regional species richness, due to much higher levels of exploitation in that geographic region. The family Testudinidae has the highest ATL (4.06) of all Testudines due to the extinction of several species of giant tortoises from Indian and Pacific Ocean islands since 1500 CE. The family Testudinidae also has an ATL higher than all other larger polytypic families (≥ 5 species) of Reptilia or Amphibia. The order Testudines is, on average, more imperiled than all other larger orders (≥ 20 species) of Reptilia, Amphibia, Mammalia, or Aves, but has percentages of CR+EN and Threatened species and an ATL (2.96) similar to those of Primates and Caudata (salamanders).

Chelonian Conservation and Biology↗

Effects of biodiversity on ecosystem functioning: a consensus of current knowledge

Humans are altering the composition of biological communities through a variety of activities that increase rates of species invasions and species extinctions, at all scales, from local to global. These changes in components of the Earth's biodiversity cause concern for ethical and aesthetic reasons, but they also have a strong potential to alter ecosystem properties and the goods and services they provide to humanity. Ecological experiments, observations, and theoretical developments show that ecosystem properties depend greatly on biodiversity in terms of the functional characteristics of organisms present in the ecosystem and the distribution and abundance of those organisms over space and time. Species effects act in concert with the effects of climate, resource availability, and disturbance regimes in influencing ecosystem properties. Human activities can modify all of the above factors; here we focus on modification of these biotic controls. The scientific community has come to a broad consensus on many aspects of the relationship between biodiversity and ecosystem functioning, including many points relevant to management of ecosystems. Further progress will require integration of knowledge about biotic and abiotic controls on ecosystem properties, how ecological communities are structured, and the forces driving species extinctions and invasions. To strengthen links to policy and management, we also need to integrate our ecological knowledge with understanding of the social and economic constraints of potential management practices. Understanding this complexity, while taking strong steps to minimize current losses of species, is necessary for responsible management of Earth's ecosystems and the diverse biota they contain. Based on our review of the scientific literature, we are certain of the following conclusions: 1) Species' functional characteristics strongly influence ecosystem properties. Functional characteristics operate in a variety of contexts, including effects of dominant species, keystone species, ecological engineers, and interactions among species (e.g., competition, facilitation, mutualism, disease, and predation). Relative abundance alone is not always a good predictor of the ecosystem-level importance of a species, as even relatively rare species (e.g., a keystone predator) can strongly influence pathways of energy and material flows. 2) Alteration of biota in ecosystems via species invasions and extinctions caused by human activities has altered ecosystem goods and services in many well-documented cases. Many of these changes are difficult, expensive, or impossible to reverse or fix with technological solutions. 3) The effects of species loss or changes in composition, and the mechanisms by which the effects manifest themselves, can differ among ecosystem properties, ecosystem types, and pathways of potential community change. 4) Some ecosystem properties are initially insensitive to species loss because (a) ecosystems may have multiple species that carry out similar functional roles, (b) some species may contribute relatively little to ecosystem properties, or (c) properties may be primarily controlled by abiotic environmental conditions. 5) More species are needed to insure a stable supply of ecosystem goods and services as spatial and temporal variability increases, which typically occurs as longer time periods and larger areas are considered. We have high confidence in the following conclusions: 1) Certain combinations of species are complementary in their patterns of resource use and can increase average rates of productivity and nutrient retention. At the same time, environmental conditions can influence the importance of complementarity in structuring communities. Identification of which and how many species act in a complementary way in complex communities is just beginning. 2) Susceptibility to invasion by exotic species is strongly influenced by species composition and, under similar environmental conditions, generally decreases with increasing species richness. However, several other factors, such as propagule pressure, disturbance regime, and resource availability also strongly influence invasion success and often override effects of species richness in comparisons across different sites or ecosystems. 3) Having a range of species that respond differently to different environmental perturbations can stabilize ecosystem process rates in response to disturbances and variation in abiotic conditions. Using practices that maintain a diversity of organisms of different functional effect and functional response types will help preserve a range of management options. Uncertainties remain and further research is necessary in the following areas: 1) Further resolution of the relationships among taxonomic diversity, functional diversity, and community structure is important for identifying mechanisms of biodiversity effects. 2) Multiple trophic levels are common to ecosystems but have been understudied in biodiversity/ecosystem functioning research. The response of ecosystem properties to varying composition and diversity of consumer organisms is much more complex than responses seen in experiments that vary only the diversity of primary producers. 3) Theoretical work on stability has outpaced experimental work, especially field research. We need long-term experiments to be able to assess temporal stability, as well as experimental perturbations to assess response to and recovery from a variety of disturbances. Design and analysis of such experiments must account for several factors that covary with species diversity. 4) Because biodiversity both responds to and influences ecosystem properties, understanding the feedbacks involved is necessary to integrate results from experimental communities with patterns seen at broader scales. Likely patterns of extinction and invasion need to be linked to different drivers of global change, the forces that structure communities, and controls on ecosystem properties for the development of effective management and conservation strategies. 5) This paper focuses primarily on terrestrial systems, with some coverage of freshwater systems, because that is where most empirical and theoretical study has focused. While the fundamental principles described here should apply to marine systems, further study of that realm is necessary. Despite some uncertainties about the mechanisms and circumstances under which diversity influences ecosystem properties, incorporating diversity effects into policy and management is essential, especially in making decisions involving large temporal and spatial scales. Sacrificing those aspects of ecosystems that are difficult or impossible to reconstruct, such as diversity, simply because we are not yet certain about the extent and mechanisms by which they affect ecosystem properties, will restrict future management options even further. It is incumbent upon ecologists to communicate this need, and the values that can derive from such a perspective, to those charged with economic and policy decision-making.

Ecological Monographs↗

Physics to fish—Understanding the factors that create and sustain native fish habitat in the San Francisco Estuary

Executive Summary The Bureau of Reclamation (Reclamation) operates the Central Valley Project (CVP), one of the nation’s largest water projects. Reclamation has an ongoing need to improve the scientific basis for adaptive management of the CVP and, by extension, joint operations with California’s State Water Project. The U.S. Geological Survey (USGS) works cooperatively with the Bureau of Reclamation to provide scientific support for the management of Reclamation’s CVP project. Major habitat restoration efforts and a new water-diversion point are planned to benefit delta smelt ( Hypomesus transpacificus ) and other species of concern while ensuring the reliability of water supply. In addition, various flow actions and management activities have been identified as possible methods to increase populations of delta smelt and salmonid ( Oncorhynchus spp.) runs of concern. The overarching goal of this cooperative project was to provide Reclamation with the scientific information needed to evaluate the efficacy of ongoing and future adaptive management actions and to improve the scientific basis for more flexible CVP operations that would achieve water-supply reliability and fish protection. The research and monitoring described in this report comprises the period 2015–19 and focuses on management issues related to native fish species of concern, especially delta smelt. Conserving the delta smelt population while providing a reliable water supply is a primary management and policy issue in California. Our approach for this cooperative project is based on the “physics to fish” concept, the idea that high-quality habitat is generated and sustained by the interaction between physical processes and the landscape. These interactions create a template for chemical and biological processes that can change across a variety of spatial and temporal scales. Following this concept, this project (hereafter referred to as “the physics to fish project”) included monitoring and studies of water flows, sediments, water quality, and invertebrate and fish dynamics across a range of spatial and temporal scales and in regions relevant to resource managers tasked with managing water supplies and ecosystem health in the San Francisco Estuary. The intent of this approach was to document the habitat conditions, important processes, and interactions among them that create high-quality habitat for native fishes so that the likely effects of future management actions (for example, habitat restoration) can be objectively assessed at the local (site-specific), regional (within subregions of the estuary), and landscape (across the entire estuary and beyond) scales. Hydrodynamically, the upper estuary (landward of Carquinez Strait) is characterized by a fixed volume of tidally exchanged water (for example, tidal prism) that interacts with the existing channel network and bathymetry to create regions with differing hydrodynamics. Our results indicate that careful study of construction or reoperation of existing infrastructure to perform management actions can help (1) improve the accuracy of hydrodynamic models; (2) further understanding of ecological effects; and (3) enhance abilities to predict ecological outcomes. At the local scale, we developed a new concept called the Lagrangian to Eulerian (LE) ratio that can be used as a tool for understanding the importance of various hydrodynamic processes in specific channels or channel networks and for forecasting transport dynamics. Channels with LE ratios<1 in a channel network or in a dead-end slough are hydrodynamically able to develop an exchange zone between two parcels of water that may have different chemical and physical properties. In a dead-end channel, there is a landward region with long residence time (no-exchange zone) and a seaward region with short residence time (high-exchange zone) that are well mixed with seaward waters. At the transition (exchange zone) between the high and no-exchange regions, a gradient will form in water-quality constituents that differ in concentration between the landward and seaward waters. Turbidity affects fish habitat and has declined through time in the San Francisco Estuary. Average turbidity across the Sacramento–San Joaquin Delta (hereafter referred to as “the Delta”) is dependent on annual hydrology. In dry years, the region around Cache Slough (known regionally as the “Cache Slough Complex”) in the northern Delta is generally more turbid than Suisun Bay and the lower Sacramento River. When the Yolo By-Pass (known regionally as “Yolo Bypass”), a large flood bypass that runs parallel to the Sacramento River in the northern Delta, is not flooding and river flows are lower, sediment is usually transported into the Cache Slough Complex because flood tides dominate ebb tides, resulting in transport of suspended sediment from seaward areas of the upper estuary into the Cache Slough Complex. These hydrodynamic conditions also favor the formation of turbidity maximums (TMs) in the Cache Slough Complex. The TMs are areas of higher suspended-sediment concentration, providing higher-turbidity habitat favored by some fishes, including delta smelt, and they can also concentrate other constituents, including phytoplankton and organic carbon that can be important in food webs. Pelagic primary production by phytoplankton is the basis for Delta food webs supporting pelagic fishes such as delta smelt; however, phytoplankton abundance in the Delta has declined during recent decades. We examined how nutrients, hydrodynamics, and other factors affect phytoplankton blooms. Based on our results, we developed three new concepts of phytoplankton bloom formation in the Delta, each associated with a distinct set of hydrologic conditions. First, productivity cascades highlighted how local processes can contribute to phytoplankton blooms observed at the regional scale. Second, we observed phytoplankton blooms in the upper San Francisco Estuary that were associated with transport out of Yolo By-Pass (transport blooms). Third, we also documented a series of phytoplankton blooms that were in the confluence area at the landward edge of Suisun Bay. The conditions leading to creation of confluence phytoplankton blooms are not yet understood, but the confluence region connects the Cache Slough Complex with Suisun Marsh. Therefore, blooms in this area have the potential to spread to large areas of the Delta. At the landscape scale, the distribution of the invasive clams ( Potamocorbula amurensis and Corbicula fluminea , hereafter referred to as “ Corbicula ”) is driven by salinity. At smaller spatial scales, the distribution of either species is sensitive to multiple factors affecting survival and reproduction, complicating efforts to predict distribution and abundance without considering local-scale conditions across the area of interest. In the Cache Slough Complex, the area landward of the exchange zone in regions with LE ratio<1 were characterized by low abundances of Corbicula probably because recruits from seaward areas are not transported past the exchange zone and because there are no landward tributaries with adult Corbicula to provide an upstream source of recruits. Corbicula biomass was highest near or downstream from the exchange zone consistent with Corbicula grazing on phytoplankton produced in the exchange zone or transported from the no-exchange zone. The severity of Corbicula grazing could be reduced by manipulating the hydrodynamic characteristics of waterways; however, the beneficial and harmful effects on the organisms meant to benefit from increased phytoplankton production, including zooplankton and fish species of concern, should be thoroughly examined before manipulating hydrodynamic characteristics. The distribution of fishes at the landscape scale is generally driven by the position of the salinity field in the estuary. The physics to fish project compared distributions of fishes at Ryer Island, a tidal wetland in Suisun Bay and a region of variable salinity, with fish distributions at the Cache Slough Complex, a freshwater region. At Ryer Island, there was an absence of freshwater invasive species and an abundance of native species, such as Sacramento splittail ( Pogonichthys macrolepidotus ), tule perch ( Hysterocarpus traskii ), and Sacramento pikeminnow ( Ptychocheilus grandis ). The native species were almost exclusively captured in wetland and nearshore shallow-water habitat regardless of water-quality conditions. In the Cache Slough Complex, our regional scale objective was to elucidate how hydrodynamic-physical habitat interactions drive fish-community structure. Our studies showed that dendritic channel systems were better able to support native species, while intertidal habitats supported those species best able to exploit the transient character of the habitat. Habitats upstream from the exchange zone were especially important in supporting high numbers of native fishes relative to within or downstream from the exchange zone. Many of the native species were associated with tidal marsh in the no-exchange zone. More pelagic-oriented, mobile species, such as Striped Bass ( Morone saxatilis ), threadfin shad ( Dorosoma petenense ), and Sacramento pikeminnow, were more affected by water-quality conditions, such as turbidity. The physics to fish concept developed in this project provides a framework for designing individual projects and for considering the cumulative effects of multiple projects in a region, using the LE ratio as a guiding metric. The physics to fish concept may also provide a suitable framework for coordinating management actions. Tidal wetlands can function in several ways in the hydrodynamic framework. Relatively small tidal wetlands with short channel networks and with LE ratios>1 are not able to maintain a landward no-exchange zone or an exchange zone. This likely means that any contributions to pelagic food webs would be limited to resources derived from wetland vegetation, which can include dissolved and particulate organic matter (detritus) and populations of consumers that can increase in abundance based on those resources. The fate of the contributed production from these channels depends on the characteristics of the receiving waters seaward of the tidal wetland. If these channels join a large system such as Suisun Bay, then any contribution is likely to be rapidly dispersed in the larger volume; however, the channel junction might provide a focal point for consumers, such as fishes, to congregate and feed on material leaving the wetland on ebb tides before it is dispersed in the larger volume. Fishes might also access these resources by entering the wetland. The physics to fish project has established a foundation and several new concepts for understanding how habitat restoration can benefit native fish populations at the local and regional levels. Many of the ideas regarding habitat restoration and channel modifications outlined in this report could help guide management actions that could improve conditions for native fishes at little or no water cost beyond water already dedicated to other management actions. A complete list of products originating from this work is provided in appendix 1.

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Fishes of Missouri River, chute, and flood plain habitats: Chapter 4 in Initial biotic survey of Lisbon Bottom, Big Muddy National Fish and Wildlife Refuge

The Lisbon Bottom Unit of the Big Muddy National Fish and Wildlife Refuge (Refuge) is approximately 2,200 acres and is the first complete unit of the Refuge. Primary objectives of the Refuge are to create and restore diverse riverine aquatic habitats and reconnect the Missouri River to its flood plain where feasible. Management seeks to accomplish these objectives by encouraging natural processes of erosion, deposition, and succession to the greatest extent possible. One of the most salient aquatic features of the Lisbon Bottom Unit is a newly created 2-mile-long free-flowing chute, or side channel (Fig. 1). This chute began forming as a levee breech scour hole during the Great Flood of 1993. The chute continued to develop during the 1995 flood and finally cut through to a flowing side channel during the 1996 flood. Extensive erosion and bank sluffing continued during 1997 due to sustained high flows that occurred throughout most of the year. The chute has progressively become wider and deeper with a developing meander pattern and channel bars have begun to form. Lisbon Bottom also contains several seasonal and permanent wetlands and is subject to periodic flooding at high Missouri River stages. Eight studies have been completed or are ongoing to evaluate Missouri River fishes associated with various habitat components of Lisbon Bottom and adjacent Missouri River reaches (Table 1). Several are part of much larger investigations to evaluate fish use of flood-created habitat features, basinwide fish assessment, and endangered or candidate species concerns. At the Lisbon Bottom Unit or in the Missouri River adjacent to the unit 54 fish species were collected (Table 2). Eight of these species have either a protected status under State or Federal laws or biologists consider them to potentially qualify for protected status. The status of the following fish is listed in Table 2: pallid sturgeon x shovelnose sturgeon hybrid, paddlefish, northern pike, sturgeon chub, sicklefin chub, ghost shiner, western silvery minnow, plains minnow, and blue sucker. Equally important to note are fish species that were not collected. The flathead chub, currently listed as State endangered and proposed for Federal listing, were not collected in any of the studies. Common fish species, collected in all seven sampling studies, included gizzard shad, common carp, river carpsucker, and freshwater drum. Paddlefish, skipjack herring, silver carp, white sucker, redfin shiner, western silvery minnow, bullhead minnow, and black bullhead were each collected in only one of the seven studies. Pflieger (1971) developed a guild system for the fishes of Missouri based on distribution patterns and centers of abundance. Fish were assigned to one of four primary faunal groups: Ozark, Big River, Lowland, and Prairie. Two secondary groups (Ozark-prairie and Ozark-lowland) were defined for species equally abundant and distributed in two of the primary areas. Two Ozark-prairie species, shorthead redhorse and white sucker, were collected on or near the Lisbon Bottom Unit. Big River species are found primarily in the Missouri and Mississippi Rivers. The environmental factors controlling fish distribution in these rivers appear to be substrate, current velocity, and turbidity. On or near the Lisbon Bottom Unit 18 Big River species were collected (Table 2). Lowland species are intolerant of siltation and high turbidity. They inhabit standing or slow-moving water with sand, fine gravel, and organic debris substrates. Two Lowland species, bullhead minnow and mosquitofish, were collected in connected scours on the Lisbon Bottom Unit. Prairie species have broader ecological tolerances than the Ozark and Lowland species. They are largely absent in high gradient streams and cool, clear waters. Prairie species make up a significant proportion of Missouri and Mississippi River fishes. Seven Prairie species were collected on or near the Lisbon Bottom Unit (Table 2). The last group, Wide-ranging, was defined as the species more widespread than the other faunal groups with broader environmental tolerances. Most Wide-ranging species occur at least occasionally in all sections of the state. Nineteen Wide-ranging species were collected on or near the Lisbon Bottom Unit (Table 2). Kubisiak (1997) found that isolated scours in the Lower Missouri River were dominated by Wide-ranging species. Connected scours also held Wideranging species but they contained Prairie species and were dominated by Big River species. Faunal group diversity and species richness were compared between isolated (Study l) and connected (Studies 4, 5, 7, and 8) scours of the Lisbon Bottom Unit. Species richness was similar with 22 species collected in isolated scours and an average of 24.8 species collected in connected scours. Isolated scour fish catches were dominated by Wide-ranging fish (59%) with 14% Big River fishes and 14% Prairie fishes. Connected scours had a larger percentage of Big River fish (32%) and fewer Wide-ranging (36%) and Prairie (13%) species than the isolated scours. In the combined studies, Wide-ranging fish dominated the connected scours. A similar comparison was made of fish from chute habitats and the Missouri River. Chute habitats include the Lisbon Bottom Chute, the chute between the right bank and island located at river mile 219, and a shallow channel running between a large sandbar complex and the channel border adjacent to Lisbon Bottom. Species richness in the chutes (35 species) was greater than that of the adjacent river (23 species). Big River species (40%) dominated the chute samples, followed by Wide-ranging at 31% Cand Prairie Cut 14%. The river samples contained equal numbers of Wide-ranging species (35%) and Big River species (35%) with 17% Prairie species. Differences in fish guild diversity and species richness may be due to numerous factors including sampling effort and sampling season. River samples were collected only in Study 3 in October; chute samples were collected in July, August, and October. Fish may have been using the chute in July and August to spawn, feed, or escape high river levels. The Lisbon Bottom Chute is a truly unique feature of the Lower Missouri River. It is the first naturally created side channel to develop since the U.S. Army Corps of Engineers tamed the river. The Corps eliminated 89% of the islands in the Missouri River between 1879 and 1954 (Funk and Robinson 1974). The chutes between the islands and the shore were shallower with less current than the main channel. These areas provided diversity to the Missouri River fish habitat, serving as nursery and feeding areas. The Lisbon Bottom Unit of the Big Muddy National Fish and Wildlife Refuge will continue to be studied in an effort to understand the complex role these areas play in river systems.

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