Title: Documentation of food-web linkages in the mainstem Columbia River – towards understanding the role of invasive species and establishing a baseline trophic state A. Abstract and statement of innovation Understanding the sources of production supporting salmonids, native and nonnative predators, white sturgeon, and other resident fish species could provide crucial information to guide management of rivers. Recovery efforts for Pacific salmon have focused on altering dam operations, improving bypass structures, and predator removal programs to increase survival of migrating juvenile salmonids. However, very little is known of how these mitigation efforts have affected the ecosystem function of the mainstem Columbia River. Further introductions of non-native and, in some instances, invasive animal and plant species have altered the trophic-level dynamics of the mainstem Columbia River. The use of naturally occurring stable isotopes to estimate the structure of the reservoir’s food web can be accomplished with minimal sampling and can provide a cost effective way to assess the current structure of the mainstem Columbia River above Bonneville Dam. Our objectives in this research are to 1) provide information to mangers that will facilitate the management of fisheries and 2) develop a unique approach to understanding how non-native species are affecting ecosystem processes and relationships in the mainstem Columbia River. In particular, how the proliferation of Eurasian watermilfoil has affected ecosystem processes. To achieve these objectives we propose to conduct a pilot study to develop methods and a sampling strategy to collect samples, determine isotopic signatures of representative trophic levels in Bonneville Reservoir, use multi-source mixing models to quantify food web sources and pathways. B. Technical and/or Scientific Background The mainstem Columbia River is an important migration corridor for endangered and threatened salmonids and also supports important fisheries for native resident fish such as white sturgeon Acipenser transmontanus. Recovery efforts for Pacific salmon have focused on altering dam operations, improving bypass structures, and predator removal programs to increase survival of seaward migrating juvenile salmonids. However, very little is known of how these mitigation efforts have affected the ecosystem function of the mainstem Columbia River. Further introductions of non-native and, in some instances, invasive animal and plant species have altered the trophic-level dynamics of the mainstem Columbia River. For instance, the invasive aquatic plant, Eurasian watermilfoil Myriophyllum spicatum L., now occupies significant portions of the littoral habitats in some mainstem reservoirs (USGS unpublished data). The current distribution of Eurasian watermilfoil has to affect the nature of primary and secondary production in these habitats and how these contribute to the trophic structure of the ecosystem. The Asian clam Corbicula fluminea and American shad Alosa sapidissima have also become abundant in the mainstem Columbia River. The Asian clam has been shown to constitute a major portion of the diet of white sturgeon (Muir et al. 1988). The abundance of American shad is likely to affect the structure of the zooplankton community. Understanding how energy is processed through food web linkages in the Columbia River will facilitate an understanding of the potential cascading effects of mitigation actions. Establishing a baseline condition will provide a means to assess the effects of mitigation or restoration actions or attempts to control or manage invasive animals or plants on the trophic level dynamics of the Columbia River ecosystem. For instance, if control efforts for Eurasian watermilfoil were conducted, what trophic levels would be most affected? Given that the Columbia River is a regulated river system, are there any potential positive effects of Eurasian watermilfoil? These types of questions will be difficult to answer without knowledge of how energy is processed through the trophic levels. The use of naturally occurring stable isotopes to estimate the structure of the reservoir’s food web can provide a cost effective way to assess the current structure of the mainstem Columbia River above Bonneville Dam. Stable isotopes provide two kinds of information for very minimal field sampling, (1) the source of an isotopic signature and (2) the change of the isotopic signature due to fractionation by physical and chemical reactions as the isotopes are consumed and metabolized in a food web (Peterson and Fry 1987). Measurements of naturally occurring stable isotope ratios of carbon (δ13C) and nitrogen (δ15N) allow us to understand the carbon flow and trophic structure over larger scales than those typically possible by experimental or observational approaches (Schell et al. 1989, Hobson 1999). Stable isotope signatures of organic materials change from food to consumer in a highly predictable manner because of selective uptake and metabolic fractionation of isotopes. Carbon isotope signature values increase typically by only 0.0 to 1.0‰ during trophic transfer (Peterson and Fry 1987), thus they can be preserved through several trophic levels and are often used as a dietary tracer (Perry et al. 2003, Bilby et al. 1998). The nitrogen isotopic signatures for consumers typically increase by 3-5‰ relative to its prey and values may increase by 10 to 15‰ in many food webs, due to successive trophic transfers (Peterson and Fry 1987). Thus, nitrogen isotopes are typically used to indicate trophic position (Vander Zanden and Rasmussen 1996, Vander Zanden et al. 1997). There is no change with sulfur at increasing trophic levels, making this a good indicator as to which plant or bacterial food sources are most important to the consumers. Furthermore, stable isotopes are temporally integrative, allowing the possibility of estimating seasonal diet composition (Fry and Sherr 1984). The use of stable isotopes can allow for the identification of food web pathways that support salmon as well as those that are affected by invasive species (e.g. shad, zebra mussels, etc). Several stable isotope studies on large river systems and reservoirs (Kendall et al. 2001; Lewis et al. 2001; Saito et al. 2001; Fry 2002; Johnson et al. 2002) have demonstrated that fish communities integrate much of the isotopic variability present at lower trophic levels. Some studies have evaluated stable isotopes in the Columbia River mainstem with a focus on evaluating suspended particulate matter that includes biota at lower trophic levels (bacteria, phytoplankton, & zooplankton) (Prahl et al. 1998, Sullivan et al. 2001). Further, studies are ongoing in the lower Columbia River and estuary (Bottom et al. 2003; BPA proposal 2000301000; see: http://www.cbfwa.org/FWProgram/ResultProposal.cfm?PPID=CE2003000030001) that have also proposed to use stable isotopes to assess the food web structure in the lower Columbia River. Similar to the assertions of Bottom et al. (2003) regarding our knowledge of food web pathways in the estuary, we suggest that there are major gaps in our understanding of how ecosystem processes have been altered. Spatial and temporal scales of connections between food webs and habitats are poorly understood in the mainstem Columbia River above Bonneville Dam. This information is important to understanding processes controlling the dynamics of local populations, communities, and ecosystems across habitats and landscapes (Polis et al. 1997, Laurance et al. 2001). Trophic connections between habitats or ecosystems may strongly influence production and structure of reservoir and stream food webs. Food webs in adjacent habitats (tributaries, reservoirs, and anthropogenic sources) and ecosystems may be strongly linked by fluxes of nutrients, detritus, or organisms (e.g. invasive species) (Finlay et al. 2002, Polis et al. 1997). By using the natural variation in δ13C and δ15N we can determine carbon sources and infer trophic positions in the reservoir ecosystem. Carbon can vary spatially and seasonally and by tracking the naturally occurring isotope variation we can also infer spatial scale of carbon flow to food webs that support mobile predators. In reservoirs, δ13C can be used to differentiate between major sources of available energy, such as, littoral production from attached algae and detritus, and pelagic production from phytoplankton, because the δ13C of the base of the littoral food web tends to be enriched in 13C (less negative than δ13C) relative to the base of the pelagic food web (France 1995). In lotic ecosystems, factors such as geomorphology and discharge (dam operations) may strongly influence carbon sources to consumers, through effects on transport of organic matter, algal production, and consumer movement. Lotic ecosystems demonstrate the spatial complexity of trophic dynamics (Finlay et al. 2002). Stable isotope techniques can provide a continuous measure of trophic position that integrates the assimilation of energy or mass flow through all the different trophic pathways leading to an organism. Stable isotopes have the potential to simultaneously capture complex interactions, including trophic omnivory, and to track energy or mass flow through ecological communities (Peterson and Fry 1987). Understanding the sources of production supporting salmonids, native and nonnative predators, white sturgeon, and other resident fish species could provide crucial information to guide management of rivers. Our objectives in this research are to 1) provide information to managers that will facilitate the management of fisheries and 2) develop a unique approach to understanding how non-native species are affecting ecosystem processes and relationships in the mainstem Columbia River. In particular, how the proliferation of Eurasian watermilfoil has affected ecosystem processes. To achieve these objectives we propose to establish a habitat based probabilistic sampling design to collect samples, determine isotopic signatures of representative trophic levels in Bonneville Reservoir, and use multi-source mixing models to quantify food web sources and pathways. C. Rationale and Significance to the Council’s Fish and Wildlife Program Specifically, the research proposed here is in support of research needs identified in the Northwest Power and Conservation Council (NPCC) Research plan. As stated previously in section B, we feel that an understanding of how energy is processed through the trophic levels of the Columbia River ecosystem will facilitate a better understanding of the effects of mitigation and restoration actions on non-target species. Establishing a baseline condition will also provide a means to assess the effects of attempts to control or manage invasive animals or plants on the trophic level dynamics of the Columbia River ecosystem. For instance, if control efforts for Eurasian watermilfoil were conducted, what trophic levels would be most affected? Given that the Columbia River is a regulated river system, are there any potential positive effects of Eurasian watermilfoil? These types of questions will be difficult to answer without knowledge of how energy is processed through the trophic levels. We feel that this research will develop methods that if fully implemented, would address the following critical management uncertainty listed in the NPCC Research Plan with respect to invasive species management (see p. 32): 1. To what extent do invasive and nonnative species affect native fish and wildlife species in the Columbia River Basin? and is consistent with the following regional research recommendations (see p. 97; Appendix H.): 9.2 Identify the interactions between native and invasive species, including: Predators, prey, food chain organisms, pathogens, and those that alter habitat structure; How competitors respond to altered systems and to restoration and recovery actions; and, How food supplies have been altered and how they can be restored. 9.3 Determine the ecological and economic consequences of invasions on native fish fauna and aquatic organisms e.g. competition, predation, and cascading trophic effects on native species, nutrient cycling, effect of management activities. Determine the impact of nonnative aquatic and terrestrial species that are invasive on salmonid recovery. Determine the impacts of nonnative species invasions i.e., rainbow and eastern brook trout, on abundance, distribution, life history and genetic diversity on bull trout, redband trout, and west slope cutthroat trout populations. 9.4 Determine the trophic impacts of nonnative species. Determine whether the economic and ecological effects of invasive species have greater impacts at some trophic levels or in specific guilds than others. Determine the current extent of the colonization of reservoirs by nonnative estuarine and mountain stream species and their role in reservoir food webs and headwater storage projects. 9.5 Determine the ecological impacts of “naturalized” non-indigenous species. 9.6 Determine whether regionally accepted non-indigenous species, such as warmwater fish, can be managed to minimize ecological effects. We propose this research as a pilot monitoring activity. Our intent is to structure the sampling design so that it can be redone in a repeatable fashion and would allow long term monitoring types of comparisons. We also propose that the sampling approach should be applicable to other mainstem reaches. As such, the research will inform regional monitoring activities such as the Pacific Aquatic Monitoring Partnership (PNAMP). D. Relationships to other projects As implied in Section’s B and C, we feel that the information that we would collect would inform a variety of projects currently being conducted in the Fish and Wildlife Program. Specifically, we feel that the information would be beneficial to projects that are charged with management and research activities for white sturgeon. We would also need to coordinate and utilize the fish sampling efforts from these projects to avoid duplication of effort and reduce the costs of this proposal. In general, we feel the information provided in this project would provide an insightful context to most management activities and research being conducted in this river reach. We will coordinate intensively with ODFW to obtain tissue and stomach samples for target fish species and other fish species opportunistically through the White Sturgeon Mitigation and Restoration in the Columbia and Snake Rivers Upstream from Bonneville Dam (BPA project 198605000) and the Northern Pikeminnow Management Program (Tom Rien personal communication). Additional samples will be obtained from the USGS white sturgeon young-of-the year index trawling that is conducted annually (BPA project 198605000). We are participating in the PNAMP (see project # 20040200, title "PNAMP Funding") technical workgroups regarding Estuaries and Large Rivers. This research will be coordinated with PNAMP to ensure consistency with regional monitoring activities and efforts to establish a universal sampling design for monitoring in the Pacific Northwest. E. Proposal objectives, work elements, and methods and monitoring and evaluation Major gaps in our understanding of how energy is processed through food webs in reservoirs on the mainstem Columbia River exist despite the importance of these habitats to popular native sport fisheries (e.g., white sturgeon) and other native resident fish species. Certain introduced animals and plants have become established and are now abundant (e.g., Asian clam, Eurasian watermilfoil, smallmouth bass, etc.). However, how these and other introduced species have affected the trophic level dynamics of these river reaches is poorly understood. Further, as part of mitigation efforts for Pacific salmon, reservoir operations are manipulated to increase the survival of juvenile salmon. Dam operations may strongly influence carbon sources to consumers, through effects on transport of organic matter, algal production, and consumer movement. Our primary biological objectives are to 1) provide information to mangers that will facilitate the management of fisheries and 2) develop a unique approach to understanding how nonnative species are affecting ecosystem processes and relationships in the mainstem Columbia River. In particular, how the proliferation of Eurasian watermilfoil has affected ecosystem processes. Bottom et al. (2001) recommended the use of natural stable isotope analyses or other methods to investigate potential food web disruptions due to habitat loss and degradation. In this research we suggest that a similar approach is warranted to understand the effects of mitigation actions for Pacific salmon and the effects of invasive species on ecosystem processes and the fish community of the mainstem Columbia River above Bonneville Dam. As per previous recommendations by the ISRP, we specifically propose to address the following hypothesis: Ho: Despite the fact that Eurasian watermilfoil is now abundant in shallow water habitats and likely constitutes a large proportion of the plant biomass in Bonneville Reservoir, the production of this aquatic plant does not contribute significantly to the food web. Management implication: Programs to limit the distribution of this invasive plant will increase the energetic inputs to Bonneville Reservoir by increasing periphyton production that will be more palatable to secondary consumers. (Note: the alternative hypothesis here is that milfoil does contribute significant resources to the food web and thus the management implication would be that the removal would alter a significant energetic pathway). We will address hypotheses 1 by determining the link between consumers and diet using the techniques outlined in Work element 2. Work element 1. Determine isotopic signatures of representative trophic levels in Bonneville Reservoir. The spatial and temporal scales of connections between food webs are poorly understood in the mainstem Columbia River above Bonneville Dam. Food webs in adjacent habitats (tributaries, reservoirs, and anthropogenic sources) and ecosystems may be strongly linked by fluxes of nutrients, detritus, or organisms (e.g. invasive species) (Finlay et al. 2002, Polis et al. 1997). In reservoirs, δ13C can be used to differentiate between major sources of available energy, such as, littoral production from attached algae and detritus, and pelagic production from phytoplankton, because the δ13C of the base of the littoral food web tends to be enriched in 13C (less negative than δ13C) relative to the base of the pelagic food web (France 1995). We will collect samples from representative habitat types that correspond to a hierarchical habitat classification system being developed by the USGS. The classification will be based on the hydrogeomorphic characteristics of reservoirs with further delineations based on channel morphology (bathymetry), shoreline type (rip-rap etc.), and other landscape and land-use characteristics (island, tributary mouths, marinas, etc.). Locations to be sampled will be randomly selected for the representative habitat classes from cells overlain as a grid on the reservoir in a GIS as per probabilistic sampling methods typically employed by EMAP. To account for temporal variability in some trophic levels, samples will be collected at different times of the year. Site selection and sampling procedures will be coordinated with regional monitoring groups such as PNAMP. Task 1.1 Riverine particulate organic matter (POM) During two periods (late spring and late fall, during periods of increased terrestrial POM input) we will obtain samples of riverine POM, separated into a phytoplankton and non-phytoplankton component for isotopic analysis. Additionally, we will obtain samples from major tributaries to Bonneville Reservoir during the two periods and during winter, high discharge events. Task 1.2 Periphyton Periphyton will be sampled by various methods. For sampling coarse/hard substrate, a stiff brush will be used to remove periphyton from a known area. A gravel sampler made of PVC may also be used depending on substrate composition. For finer substrate, the inverted petri dish method outlined in the NAWQA protocol will be utilized. Epiphytic algae will be sampled in conjunction with aquatic macrophytes described below. Task 1.3 Aquatic macrophytes Macrophyte samples will be collected using a stovepipe sampler. Specific collection locations will be determined using a point-intercept (grid) or line-intercept (transect) method described by Madsen (1999). Macrophytes will be rinsed with water, brushed or agitated for sampling attached algae. Task 1.4 Zooplankton and benthos Riverine zooplankton will be collected by plankton net sampling at locations that correspond to a habitat classification system that is being developed for Bonneville Reservoir by the USGS. Vertically integrated zooplankton samples will be taken with 2 Wisconsin-style plankton nets attached side-by-side to a metal frame. One net will have a mesh size of 64 μm to capture cladocerans and postnaupliar copepods, and the other net will have a mesh size of 35 μm to capture rotifers and nauplii. The nets will be towed from near the bottom to the surface, obliquely in flowing waters, and vertically in low flow and slack water areas. Zooplankton taxa may be composited into a single sample for isotope analysis. Benthic organisms will be collected using a ponar dredge, disturbance methods, or benthic cores. To collect invertebrate taxa on aquatic macrophytes we will use a stovepipe sampler. Crayfish spp. will be collected using traps. Taxa will be sampled and classified according to the following functional (trophic) groups: 1) pelagic zooplankton, 2) benthic infauna, separated into representative suspension feeders and detritivores; certain amphipod taxa, bivalves, crayfish spp., etc. 3) epibenthic organisms, e.g., mysids 4) aquatic insects, e.g., chironomid larvae and pupae 5) fallout insects, separated into those of aquatic and terrestrial origin e.g., adult dipterans Task 1.5 Fish We will coordinate intensively with ODFW to obtain tissue and stomach samples for target fish species and other fish species opportunistically through the White Sturgeon Mitigation and Restoration in the Columbia and Snake Rivers Upstream from Bonneville Dam (BPA project 198605000) and the Northern Pikeminnow Management Program (Tom Rien personal communication). Additional samples will be obtained from the USGS white sturgeon young-of-the year index trawling that is conducted annually (BPA project 198605000). Specifically we will obtain at least five tissue samples from multiple year classes of white sturgeon Acipenser transmontanus, Common carp Cyprinius carpio, Northern pikeminnow Ptychocheilus oregonensis , smallmouth bass Micropterus dolomieu, and American shad Alosa sapidissima. Other common species of fish will also be collected and processed. Dorsal muscle samples from fish larger than 67 mm total length will be collected. For smaller fish (less than 67 mm), whole fish will be collected to assure a large enough sample (Saito et al. 2001). Stomach samples will also be taken, preserved and processed in the laboratory to corroborate what fish are eating. Since ODFW are already processing stomach samples as part of the Northern Pikeminnow Management Program, we will coordinate with them to ensure that are methodologies are consistent and that we minimize duplication of effort so that we are as cost effective as possible. Task 1.6 Sample processing for isotopes All samples will be dried for at least 48 h in a 60º C oven and then ground to a homogenous powder using a mortar and pestle. To economize on mass spectrometer costs, individual fish samples can be pooled into composite samples according to length classes that approximate age classes and reservoir locations. Composite samples should contain an equal mass of each individual sample (Saito et al 2001). In order to characterize the mean stable isotope and the associated variation about the mean for each taxonomic grouping across time periods and locations, a minimum of five samples will be processed (Lancaster and Waldron 2001). A 1-2 mg sample will be individually processed for δ13C, δ15N, and δ34S signatures in a mass spectrometer. The quantity measured in stable isotope analysis, δ, is the relative difference between isotope ratios of the sample and a standard: δ = Rsample - Rstandard/ Rstandard • 103 where δ (‰) is the per mil difference, Rsample is the isotopic ratio of the sample, and Rstandard is the isotopic ratio of the standard. The standards will we employ are Vienna Pee Dee Belmnite for 13C/12C and atmospheric N2 for 15N/14N (Lajtha and Michener 1994). Isotopic δ34S signatures will be analyzed using similar methods. Stable isotope samples will be sent to the Colorado Plateau Stable Isotope Laboratory based at Northern Arizona University in Flagstaff, AZ for analysis. The laboratory processes isotope samples for a diverse array of research organizations (see: http://www4.nau.edu/cpsil/). Work element 2. Establish food web linkages. Task 2.1 Estimate autotrophic sources for consumers. Simple mixing models are limited by the number of sources and tracers used, therefore they do not adequately describe complex food webs having multiple contributing sources and pathways (Lubetkin and Simenstad 2004). We propose to use a similar approach as that of Lubetkin and Simenstad (2004) that use companion models SOURCE and STEP that overcome these limitations and can model complex food webs in aquatic ecosystems. The SOURCE model estimates the assortment of autotrophic sources for consumers using the equations: s1 s2 s3 ··· sn 1 (a) s1t1 s2t2 s3t3 ··· sntn L Ct (b) s1u1 s2u2 s3u3 ··· snun L Cu (c) s1v1 s2v2 s3v3 ··· snvn L Cv (d) where t,u, and v are tracers; represent average trophic level shifts; Ct, Cu, Cv, represent overall consumer tracer levels, which reflect it’s diet at trophic level L; and 0 si 1. An exact solution for the above equations cannot be calculated, therefore, sets of linear equations for subsets of sources are solved iteratively to model the mixture of multiple sources. Task 2.2 Determine direct links between consumers and their diets The STEP model is used to determine links between consumers and their diets. This model estimates the isotopic signature for each consumer as a composite of its diet modified by one metabolic fractionation, and is represented by the equations below: f1 f2 f3 ··· fm 1 (e) f1t1 f2t2 f3t3 ··· fmtm Ct f1u1 f2u2 f3u3 ··· fmum Cu (f) (g) f1v1 f2v2 f3v3 ··· fmvm Cv (h) where f are fractions that m potential foods contribute to a consumers diet. The STEP model estimates direct diets of each consumer and calculates the trophic level position using the equation: f1L1 f2L2 f3L3 ··· fmLm 1 L (i) where, fi is the fraction each food source contributes to the diet of the consumer, and Li is the trophic level for each food source, and L is the trophic level of interest. For a more complete and thorough description of the model and how mathematical solutions are determined, see Lubetkin and Simenstad (2004). Task 2.3 Validate food web model Model validation will be done in a similar manner to Lubetkin and Simenstad (2004) where simulations will be run on data assumed to be perfect (no variability) and data assumed to be more representative of multiple samples (introduced variability). By performing these simulations we can evaluate how the model performs under ideal conditions, focusing on the mathematical validity of the model and how natural variability in the data would affect the model’s performances. Work element 3. Report results in technical reports, peer reviewed publications, and presentations. All work will be summarized and presented in a final technical report, peer reviewed publications, and presentations to professional and public interest groups. F. Facilities and Equipment The U.S. Geological Survey (USGS) is the primary research arm of the Department of the Interior, which manages nearly one-fifth of all land in the United States. The Department of the Interior co-chairs the National Invasive Species Council, and the USGS plays a significant role in implementing the National Invasive Species Management Plan, called for in the Presidential Executive Order 13112 on Invasive Species. To meet the goals of the Plan, the USGS Invasive Species Program provides management-oriented research and delivers information needed to prevent, detect, control, and eradicate invasive species, and to restore impaired ecosystems. The Columbia River Research Laboratory has been conducting research in the Columbia River Basin for over 25 years. The laboratory has state of the art computing resources and GIS capabilities, and approximately 30 research vessels deployed throughout the basin. In addition to the facilities, equipment, and staff specifically at the Columbia River Research Laboratory, we will seek and utilize the expertise that exists in other centers, disciplines, and programs in the USGS. G. Literature cited Bilby, R. E., Fransen, B. R., Bisson, P. A., & Walter, J. K. 1998. Response of juvenile coho salmon (oncorhynchus kisutch) and steelhead (oncorhynchus mykiss) to the addition of salmon carcasses to two streams in southwestern washington, U.S.A. Canadian Journal of Fisheries and Aquatic Sciences, 55(8), 1909-1918. Bottom, D. L., C. A. Simenstad, A. M. Baptista, D. A. Jay, J. Burke, K. K. Jones, E.Casillas, and M. H. Schiewe. 2001. Salmon at River’s End: The Role of the Estuaryin the Decline and Recovery of Columbia River Salmon. U.S. National MarineFisheries Service, Seattle. 271 pg. Bottom et al. 2003. Historic Habitat Opportunities and Food-Web Linkages of Juvenile Salmon in the Columbia River Estuary and Their Implications for Managing River Flows and Restoring Estuarine Habitat, Bonneville Power Administration FY 2003 Provincial Project Review. Finlay, J. C., Khandwala, S., & Power, M. E. 2002. Spatial scales of carbon flow in a river food web. Ecology, 83(7), 1845-1859. France, R. L. 1995. Differentiation between littoral and pelagic food webs in lakes using stable carbon isotopes. Limnology and Oceanography, 40(7), 1310-1313. Fry, B. 2002. Conservative mixing of stable isotopes across estuarine salinity gradients: A conceptual framework for monitoring watershed influences on downstream fisheries production. Estuaries, 25(2), 264-271. Fry, B. and E. B. Sherr. 1984. δ13C measurements as indicators of carbon flow in marine and fresh-water ecosystems. Contributions in Marine Science 27:13-47. Hobson, K. A. 1999. Tracing origins and migration of wildlife using stable isotopes: a review. Oecologia 120:314-326. Johnson, B. M., Martinez, P. J., & Stockwell, J. D. 2002. Tracking trophic interactions in coldwater reservoirs using naturally occurring stable isotopes. Transactions of the American Fisheries Society, 131(1), 1-13. Kendall, C., Silva, S. R., & Kelly, V. J. 2001. Carbon and nitrogen isotopic compositions of particulate organic matter in four large river systems across the united states. Hydrological Processes, 15(7), 1301-1346. Lajtha, K. and R. H. Michener, editors. 1994. Stable isotopes in ecology and environmental science. Oxford, UK, Blackwell Scientific Publications. Lancaster, J. and S. Waldron. 2001. Stable isotope values of lotic invertebrates: Sources of variation, experimental design, and statistical interpretation. 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Diet of juvenile and subadult white sturgeon in the lower Columbia River and its estuary. Calif. Fish Game 74:49-54. Perry, R. W., Bradford, M. J., & Grout, J. A. 2003. Effects of disturbance on contribution of energy sources to growth of juvenile chinook salmon (oncorhynchus tshawytscha) in boreal streams. Canadian Journal of Fisheries and Aquatic Sciences, 60(4), 390-400. Peterson, B. J., and B. Fry. 1987. Stable isotopes in ecosystem studies. Annual Review of Ecology and Systematics 18:293-320. Polis, G. A., W.B. Anderson, & R. D. Holt. 1997. Toward an integration of landscape and food web ecology: the dynamics of spatially subsidized food webs. Annual Review of Ecological Systems 28:289-316. Prahl, F.G., L.F. Small, B.A. Sullivan, J. Cordell, C.A. Simenstad, B.C. Crump, & J.A. Baross. 1998. Biogeochemical gradients in the lower Columbia River. Hydrobiologia 361:37-52. Saito, L., Johnson, B. M., Bartholow, J., & Hanna, R. B. 2001. Assessing ecosystem effects of reservoir operations using food web-energy transfer and water quality models. Ecosystems, 4(2), 105-125. Schell, D. M., Saupe, S. M., & Haubenstock, N. 1989. Bowhead whale (balaena mysticetus ) growth and feeding as estimated by delta super(13)C techniques. Marine biology.Berlin, Heidelberg, 103(4), 433-443. Sullivan, B. E., Prahl, F. G., Small, L. F., & Covert, P. A. 2001. Seasonality of phytoplankton production in the columbia river: A natural or anthropogenic pattern? Geochimica et Cosmochimica Acta, 65(7), 1125-1139. Vander Zanden, M. J., & Rasmussen, J. B. 1996. A trophic position model of pelagic food webs: Impact on contaminant bioaccumulation in lake trout. Ecological Monographs, 66(4), 451-477. Vander Zanden, M. J., Cabana, G., & Rasmussen, J. B. 1997. Comparing trophic position of freshwater fish calculated using stable nitrogen isotope ratios ( delta super(15)N) and literature dietary data. Canadian Journal of Fisheries and Aquatic Sciences, 54(5), 11421158. H. Key personnel Timothy D. Counihan U.S. Geological Survey Biological Resources Division Columbia River Research Laboratory 5501-A Cook Underwood Road Cook, Washington 98605 Education: M. S., Wildlife Science, New Mexico State University, 1991 B. S., Biology, Montana State University, 1989 Work Experience 1993-Present Research Fishery Biologist, U.S. Geological Survey, Biological Resources Division, Columbia River Research Laboratory, Cook, WA. Current responsibilities: Current projects include research examining the distribution and abundance of Eurasian watermilfoil in relation to the hydraulic characteristics of Bonneville Reservoir, Columbia River with emphasis on the effects of hydrosystem operation on the distribution of this invasive aquatic plant and modeling the potential spread of New Zealand mudsnail in Bonneville Reservoir. Other research includes modeling the survival of endangered and threatened juvenile salmonids as they pass through the lower Columbia River dams and working on a multi-agency team to formulate a habitat classification system and habitat and ecological monitoring program for the lower Columbia River. Expertise: Primary areas of expertise include the design and analysis of fish and ichthyoplankton surveys, survival assessment using mark-recapture models, white sturgeon ecology, larval fish ecology, and biometrics. Publications and Reports Counihan, T. D. J.M. Hardiman and J.R. Hatten. In preparation. Modelling the influence of channel morphology and reservoir operations on the distribution of Eurasian watermilfoil in an impoundment of the Columbia River. Simenstad, C.A., J.L. Burke, I. R. Waite, T.D. Counihan, and J.R. Hatten. In preparation. Lower Columbia River and Estuary Ecosystem Classification: Phase I. Prepared for Lower Columbia River Estuary Partnership Counihan, T. D., A. I. Miller, and M. J. Parsley. 1999. Indexing the relative abundance of age-0 white sturgeon in an impoundment of the lower Columbia River from highly skewed trawling data. North American Journal of Fisheries Management. Counihan, T. D., and C. N. Frost. 1999. Influence of externally attached transmitters on the swimming performance of juvenile white sturgeon. Transactions of the American Fisheries Society. Counihan, T. D., A. I. Miller, M. G. Mesa, and M. J. Parsely. 1998. The effects of dissolved gas supersaturation on white sturgeon larvae. Transactions of the American Fisheries Society 127:316-322. Jill M. Hardiman U.S. Geological Survey Biological Resources Division Columbia River Research Laboratory 5501-A Cook Underwood Road Cook, Washington 98605 Education Colorado State University, M.S. Fishery and Wildlife Biology, 2003 University of Puget Sound, B.S. in Biology, 1992 Work Experience Fishery Biologist - USGS/Columbia River Research Laboratory, Cook, WA -3/04 to present Prepare professional presentations, annual reports, research proposals, and manuscripts Evaluate juvenile salmonid passage and survival at three hydroelectric facilities on the Lower Columbia River Investigate hydrodynamic modeling and community dynamics with regard to invasive species in Bonneville Dam reservoir Fishery Biologist IV - Johnson Controls Inc. contract for USGS/Columbia River Research Laboratory - 8/03-3/04 Prepare professional presentations, annual reports, research proposals, and manuscripts Use SAS to analyze and summarize age frequency data from 14 years of white sturgeon trawl surveys Prepare Geographic Information Systems (GIS) maps for study design, sampling protocols, presentations, spatial analysis, and reports for Eurasian Watermilfoil surveys Graduate Teaching Assistant/Research Assistant - Colorado State University, Department of Fishery and Wildlife Biology, Fort Collins, CO 8/00-5/03 Conduct limnology surveys for stratified zooplankton tows, temperature, dissolved oxygen and light profiles, conductivity, and Secchi depth measurements Track zooplankton seasonal densities to determine effects on age-0 kokanee distribution and use in bioenergetics models to determine stomach capacity, feeding rates, and time to satiation Conduct sampling of age-0 kokanee with hydroacoustics, gill nets, seines and trawls in rivers and reservoirs Collect benthic, zooplankton, and fish samples for stable isotope analysis Selected Publications Counihan, T. D., J. M. Hardiman, C. E. Walker, A. L. Puls, and G. S. Holmberg. 2005. Survival estimates of migrant juvenile salmonids in the Columbia River through Bonneville Dam using radio-telemetry, 2005. Prepared by the U. S. Geological Survey, Cook, WA for the U. S. Army Corps of Engineers, Portland, OR. Hardiman, J. M., B. M. Johnson, and P. J. Martinez. 2004. Do predators influence distribution of age-0 kokanee in a Colorado reservoir? Transactions of the American Fisheries Society 133:1366-1378. Johnson, B. M. and J. M. Hardiman. 2003. Kokanee in Blue Mesa Reservoir: Causes for their decline and strategies for recovery, Final Report. Fisheries Ecology Laboratory, Department of Fishery and Wildlife Biology, Colorado State University. Prepared for: National Park Service, Gunnison, CO, November 3, 2003. Christopher E. Walker U.S. Geological Survey Biological Resources Division Columbia River Research Laboratory 5501-A Cook Underwood Road Cook, Washington 98605 Education Portland State University, M.S. Environmental Sciences/Biology Portland State University, B.S. Environmental Science/Biology Work Experience United States Geological Survey, BRD, Cook, WA, 5/01-present, Biologist Duties: report and proposal writing, publication writing and review, prepare graphs and charts for reports and presentations, collection and analysis of other biological data, experimental set-up and study design, database management and use of GIS, statistical analysis, analyze samples for chlorophyll a, ID and enumerate phytoplankton, coordinate project with contractors and other agencies Portland State University, Portland, OR, 8/99-4/01, Research Assistant Duties: collecting water samples, macroinvertebrates, and algae, mini-piezometer installation (small wells), took water and biological samples, measured stream habitat parameters (i.e. discharge, morphometry, riparian conditons, instream substrate), analyzed samples for nutrients (ortho-phosphate, NO3+NO2, NH4+, TP, TN), chlorophyll a, ash-free dry mass, identify aquatic macroinvertebrates and algae, statistical analysis using multivariate and univariate approaches for relating biotic communities to environmental variables, report writing, publication writing and review, methods development, prepared graphs and charts for reports and presentations, experimental setup, data base management, spatial analyses using GIS Department of Environmental Quality, State of Oregon, 6/00-9/00 Field Tech. Duties: conduct water quality, habitat, macroinvertebrate, and fish, included backpack electroshocking, use of other instruments such as GPS, conductivity meter, pH meter, performing DO titrations and Fish measurements/Ids (salmonids) and Macroinvertebrate IDs, stream measurements such as discharge, measurements of riparian conditions, and instream substrate Selected Publications Walker, C.E., Schrock, R.M., Reilly, T.J., and A.L. Baehr. 2005. A direct immunoassay for detecting diatoms in groundwater as an indicator of the direct influence of surface water. Journal of Applied Phycology 17(1):81-90. Walker, C.E., and Y. Pan. 2005. Changes in Diatom Assemblages Along an Urban-to-Rural Gradient. Hydrobiologia. (in press) Sonoda, K. Yeakley, J.A. Walker, C.E. 2001. Near-stream landuse effects on streamwater nutrient distribution in an urbanizing watershed. Journal of the American Water Resources Association 37(6):1517-1532. Counihan, T.D., Holmberg, G.S., Walker, C.E., and J.M. Hardiman. 2003. Survival Estimates of migrant Juvenile Salmonids in the Columbia River through John Day Dam using RadioTelemetry, 2003. Prepared by U. S. Geological Survey, Cook, Washington for the U.S. Army Corps of Engineers, Portland, Oregon, contract W66QKZ31005684. Amy L. Puls U.S. Geological Survey Biological Resources Division Columbia River Research Laboratory 5501-A Cook Underwood Road Cook, Washington 98605 Education M.S., Marine Biology, Oregon Institute of Marine Biology, University of Oregon, 2002 B.S., Biology, University of Oregon, 1997 Work Experience Fisheries Biologist - USGS, Columbia River Research Laboratory, Cook, WA, 4/04-present Investigated survival and egress of juvenile salmonids during hydroelectric dam passage on the Columbia River using radio telemetry. Design and plan experiments examining the behavior and survival of radio-tagged juvenile salmonids at hydroelectric projects Use SAS to manage, manipulate, and analyze large datasets of fish distribution data Compose and prepare research proposals, technical presentations, reports and manuscripts Formulate and implement quality assurance/quality control procedures for data processing Graduate Research; Zooplankton Ecology - Oregon Institute of Marine Biology, Charleston, OR, 1/98 - 3/02 Investigated spatio-temporal variation of abundance for ~20 zooplankton species in relation to physical forcings and larval behavior in the Coos Bay estuary, Oregon. Gained insights into zooplankton transport mechanisms and exchange of zooplankton between the estuary and the ocean. Developed hypotheses, designed study, and wrote grant proposals Collected biological data from boats using neuston nets and standard plankton nets; collected physical data using CTD, DO meter, velocity meter, and GPS 2000+ hours spent identifying and enumerating zooplankton Research Assistant - Oregon Institute of Marine Biology, Charleston, OR, 1/98 - 12/00 Investigated physical and biological processes influencing the larval transport and recruitment of Dungeness crabs and other marine invertebrates as part of the Pacific Northwest Coastal Ecosystems Regional Study (PNCERS), a multi-institute, interdisciplinary study. Collected and preserved 3-5 light trap samples daily Constructed and maintained light traps Used dissecting scope to identify and enumerate larvae from samples Select Publications Puls, A.L. 2002. Transport of zooplankton in South Slough, Oregon. Master's thesis. 96p. University of Oregon,Eugene. Puls, A.L. 2001. Arthropoda: Decapoda. Pages 89-161 in A.L. Shanks, editor. An Identification Guide to the Larval Marine Invertebrates of the Pacific Northwest. Oregon State University Press, Corvallis. Meeuwig, M.H., AL Puls, J.M. Bayer. Provisionally accepted. Survival and tag retention of Pacific lamprey larvae and macrophthalmia marked with coded wire tags. North American Journal of Fisheries Management. Puls, A.L., G.C. Roegner, A.L. Shanks. In prep. Fluxes of invertebrate larvae through South Slough, Oregon.