Proposal: Tahoe Science Program Round 12 Request for Proposals I. Title Page Project Title: Subtheme addressed by this proposal Principal Investigator and Receiving Institution Co-Principal Investigator Co-Principal Investigator Agency Collaborator Agency Collaborator Agency Collaborator Agency Collaborator Grants Contact Person Funding requested: Total cost share (value of financial and in-kind contributions): Understanding the decline of deepwater sensitive species in Lake Tahoe: What is responsible, eutrophication or species invasions? 2c: Increasing our understanding of special status species and communities Dr. Sudeep Chandra Board of Regents, NSHE, obo University of Nevada- Reno Department of Natural Resources and Environmental Science 1000 Valley Road/MS 186, Reno, Nevada 89512 Phone: 775-784-6221, Fax: 775-784-4530 Email: sudeep@cabnr.unr.edu Dr. John Reuter University of California- Davis UC Davis Tahoe Environmental Research Center, Davis, CA 95617 Phone: 530-304-1473, Fax: 530-754-9364 Email: jereuter@ucdavis.edu Dr. Eliska Rejmankova University of California- Davis Dept of Environmental Science and Policy University of California, Davis, Davis, CA 95617 Phone: 530-304-1473, Fax: 530-754-9364 Email: erejmankova@ucdavis.edu Shane Romsos and Patrick Stone Tahoe Regional Planning Agency PO Box 5310, Stateline, NV 89449 Phone: 775-589-5201, 775-589-5213, Fax: 775-588-4527 Email: sromsos@trpa.org, pstone@trpa.org Kevin Thomas California Department of Fish and Game North Central Region 1701 Nimbus Road, Rancho Cordova, CA 95670 Phone: 916-358-2845, Fax: 916-358-2912 Email: kthomas@dfg.ca.gov Patrick Wright California Tahoe Conservancy 1061 Third Street , South Lake Tahoe, CA 96150 Phone: 530-543-6002, Fax: 530-542-5591 Email: pwright@tahoe.ca.gov Elizabeth Harrison Nevada Division of State Lands 901 S Stewart Street, Suite 5003, Carson City, NV 89701 Phone: 775- 684-2736, Fax: 775-684-2721 Email: eharrison@lands.nv.gov Carla Beier University of Nevada- Reno Office of Sponsored Projects/MS 325, Reno, NV 89557 Phone: 775-784-6754, Fax: 775-784-6680 Email: ospadmin@unr.edu $ 281,623 $0 Proposal: Tahoe Science Program Round 12 Request for Proposals II. Proposal Narrative a. Project abstract A comparison of historical and contemporary benthic surveys showed a large decline in bottom invertebrate and plant occurrence in the deepwater environment of Lake Tahoe (Caires et al. 2010). The decline in plants may be directly related to reductions in water clarity over the past 4 decades, while reductions in the invertebrate community may be related to plant declines and invasive species introductions. Two of Lake Tahoe’s unique bottom species (a blind amphipod and a deep-water stonefly), found nowhere else in the world, are at high risk given their significant reduction since the 1960s. Currently, the deepwater benthic environment in Lake Tahoe is not being monitored and is not well understood. We will examine the spatial distribution of these deepwater special status plant and invertebrate communities and, in doing so, gather information about their biology and ecology. Specifically, we will study the life cycles of special status plants and invertebrates and feeding strategies of endemic invertebrates. We will also measure native plant photosynthetic rates at varying light conditions and relate observed plant responses to long term subsurface irradiance data in order to understand the depths at which light has been limiting for deepwater plants over the last four decades and how this has affected changes in their vertical distribution. The influence of future clarity improvement (as part of the TMDL strategy) on the redistribution of this special deepwater plant community will be evaluated using a model to be developed for this purpose. Non-native invertebrate effects on special status communities will also be analyzed in the laboratory. We hope to determine mechanisms that have contributed to the decline of these unique deepwater plant and invertebrate communities and develop appropriate restoration strategies. A monitoring plan will be developed to allow managers to track special status community response to restoration strategies, such as changes in light penetration and/or reductions in non-native species. Monitoring of relatively long-lived organisms such as aquatic plants and invertebrates provides an important biological indicator of the overall health of the system, one that has received little attention in Lake Tahoe. b. Justification statement Research conducted by our team in 2008-2010 suggests that benthic plant and invertebrate distribution and density have been substantially reduced in Lake Tahoe since the 1960s. Our proposal explicitly addresses subtheme 2c, “Increasing our understanding of special status species and communities,” by determining mechanisms leading to the decline of the endemic Tahoe stonefly, blind amphipod, and associated deepwater plant communities. These special status communities can be used to assess alterations and trends that have occurred in pelagic (through coupling to the bottom) and benthic environments. We will study these communities and their role in ecological processes through field collections and laboratory experiments, along with a model to help us understand how these communities have responded to decreased clarity in the lake. Based on our findings, we will make recommendations for actions that could improve habitat and potentially restore these special status communities. This project is highly relevant to current efforts in the Tahoe Basin to develop a complete set of environmental indicators that evaluate lake condition and compliment the open water indicators currently employed to quantify changes to the lake. c. Concise background and problem statement Deepwater special status communities in Lake Tahoe. Historical invertebrate surveys from the benthic environment in Lake Tahoe have been extremely limited; however, they do reveal the existence of a variety of unique, endemic benthic invertebrate taxa, mostly found in deep water (Frantz and Cordone 1966, 1996). Lake Tahoe’s endemic benthic faunal assemblage includes two species of blind amphipod (Stygobromus spp.), typically found in subterranean environments, and the Tahoe stonefly (Capnia lacustra) that is one of only two taxa in the world known to live its entire life-cycle underwater (Baumann 1984). Lake Tahoe also has a unique assemblage of deepwater plant beds, including several species of stoneworts, liverworts, and mosses, concentrated historically at depths around 100 meters (Frantz and Cordone 1967). Recently we synthesized historical Proposal: Tahoe Science Program Round 12 Request for Proposals information and compared it to contemporary surveys of benthic invertebrate communities conducted in 2008-09. Our findings indicate that there has been a severe (75%) decline in lakewide, weighted benthic invertebrate densities since the 1960s, with blind amphipods and Tahoe stoneflies showing the greatest declines at >99% and 94%, respectively (Caires et al. 2010; Figures 1a-1c). A comparison of contemporary and historical benthic samples also shows a reduction in the occurrence of aquatic plants in benthic samples, and a shift of plants to shallower waters (Figure 1d). Despite this severe decline in deepwater plant and invertebrate abundance in Lake Tahoe over the past 50 years, the biology, ecology and current distribution of these organisms is not well understood. Benthic invertebrate densities were highest in plant-dominated samples in the 1960s collections (Figure 2), suggesting that deepwater plant communities are important to native invertebrate assemblages. The association between the endemic Tahoe stonefly and deepwater plants in Lake Tahoe has been described (Jewett 1965), and the Tahoe stonefly was located in plant beds in the South Lake Tahoe area in recent surveys. Conversely, there is no clear association between aquatic plants and blind amphipods in Lake Tahoe. In fact, in the 1960s, blind amphipods were most abundant in areas that were deeper than the depth limits of most aquatic plants in the lake (Frantz and Cordone 1996). Although the dietary preferences and requirements of blind amphipod species in Lake Tahoe have not been studied, many other amphipod taxa rely on organic matter and associated bacterial assemblages for food (Covich and Thorp 2001). It is possible that blind amphipods in Lake Tahoe are dependent on detritus and associated bacteria largely derived from these deepwater plant beds. Information about the life history and feeding habits of the Tahoe stonefly and the blind amphipod could provide insight into associations between these special status invertebrates and their preferred habitat. Our surveys in 2008-09 revealed two locations in which Tahoe stoneflies were located: McKinney Bay and South Lake Tahoe. Deepwater plant beds were also located in South Lake Tahoe and blind amphipods were found in McKinney Bay. Dispersed sampling in these “hotspot” areas where special status communities have been found previously will provide information on their finer scale spatial distribution in these areas. Once target areas are identified, focused, frequent sampling will provide insight into the life histories of special status taxa, as determined by seasonal developmental state. Stable isotope analysis will allow for determination of feeding habits of endemic invertebrates. Carbon and nitrogen stable isotopes can show energy flow in food webs, where δ15N signatures indicate the trophic position of a target consumer and δ13C signatures can be used to discriminate between benthic and pelagic sources of energy in a consumer (Vander Zanden et al. 2003). Determination of feeding position and food sources will help link endemic invertebrates to their preferred habitat and may provide some insight as to the source of their observed declines. Alterations to the clarity and benthic ecology of Lake Tahoe in the last 50 years. Increased nutrient and sediment inputs to Lake Tahoe since the 1960s have caused primary productivity to steadily increase while clarity has steadily decreased (Goldman 1988, Jassby et al. 2003, TERC 2011; Figure 3). Efforts to improve lake clarity and water quality through environmental thresholds established by the Tahoe Regional Planning Agency (TRPA) and the development of a Total Maximum Daily Load (TMDL) by the Lahontan Water Board and Nevada DEP are intended to help reverse these trends. Measurements of physical, chemical, and biological parameters in the lake, such as transparency, nutrient content, and phytoplankton primary productivity/chlorophyll have been useful as indicators of changes in Lake Tahoe’s water quality (Goldman 1988; Jassby et al. 1994, 2003; Swift et al. 2006). Current monitoring efforts provide snapshots of the open water lake condition. Chandra et al. (2005) suggest that observed changes to the open-water of Lake Tahoe are directly impacting bottom processes. Thus, developing a method to assess changes at the bottom of the lake will integrate open-water monitoring with longer-term change to benthic habitats. Benthic plant distribution is regulated by underwater light availability (e.g. Schwarz et al. 1996, 1999, Proposal: Tahoe Science Program Round 12 Request for Proposals 2000), and increases in open water primary production and sediment particles in lakes can result in shading of benthic environments (Sand-Jensen and Borum 1991). The predictable growth responses of aquatic plants to changes in subsurface light availability suggest their potential as direct responders to water clarity changes. Decreased clarity in Lake Tahoe has decreased subsurface irradiance, likely contributing to the shift of deepwater plant communities to shallower water. Deepwater plants in Lake Tahoe also have shallow water depth limits due to growth limitations caused by wave action and unsuitable substrate (Frantz and Cordone 1967). Thus, there has been a narrowing depth band of suitable habitat for deepwater plants in Lake Tahoe. Despite apparent declines in overall density, deepwater plants were located in recent benthic samples, suggesting that an improvement in water clarity could allow plant communities to reestablish at deeper depths. In other lakes, benthic plants have readily responded to changes in lake clarity (Zhu et al. 2006, Schwarz et al. 1999). For example, in a large oligotrophic New Zealand lake, Chara responded negatively to a decline in lake clarity, but recovered within two years when clarity improved (Schwarz et al. 1999). Photosynthesis-irradiance (P-I) curves describe plant responses to changing light regimes, and they indicate light-compensation points below which there is insufficient light to compensate for respiratory losses. The development of P-I curves for the most common deepwater plant taxa in Lake Tahoe would provide an indication of the growth limitations of special status plant communities. This, combined with subsurface irradiance data that has been collected in Lake Tahoe since the late 1960s, could indicate the depth limit of aquatic plant growth over the past 40+ years. Similarly, the vertical extent plant growth and abundance could be projected into the future based on efforts to improve clarity. It is possible that non-native species have also negatively affected special status communities in Lake Tahoe over the past four decades. In 1963-65, shortly after the 1962-63 benthic invertebrate survey, 333,000 mysid shrimp (Mysis relicta) were introduced to Lake Tahoe as a food source for lake trout (Salvelinus namaycush) (Linn and Frantz 1965, Frantz and Cordone 1996). The mysid shrimp established a large population (approximately 300 per m2) by 1971 and subsequently caused the disappearance of two cladoceran species (Richards et al. 1975, Goldman et al. 1979, Threlkeld 1981). Mysid shrimp in Lake Tahoe feed at night in the water column and migrate to the bottom of the lake during the day to avoid fish predation, for a total daily vertical migration of up to 1000 m (Rybock 1978). These omnivorous crustaceans consume a variety of benthic and pelagic food items, including zooplankton, amphipods, and even small fish larvae (Parker 1980, Sealer and Binowski 1988, Johannsson et al. 2001, Wilhelm et al. 2002, Bailey et al. 2006). Mysid shrimp appear to be extremely abundant on sediment surfaces during the day in South Lake Tahoe, as seen in video surveys taken by a remotely operated vehicle in 1990 (see Beauchamp et al. 1992). It is possible that mysid shrimp have taken advantage of endemic benthic invertebrates in Lake Tahoe as a food source, especially exposed blind amphipods in profundal areas too deep for plant growth. Another omnivorous, non-native invertebrate, the signal crayfish (Pacifastacus leniusculus) may also be contributing to the decline of native plant and invertebrate species. Although present in the lake since 1895, crayfish numbers have nearly doubled since the 1960s (Chandra 2011; Figure 4). Adult crayfish in Lake Tahoe target benthic periphyton and plants for food, while juvenile crayfish consume mainly small benthic macroinvertebrates (Flint 1975). Mysid shrimp in Lake Tahoe have not been surveyed recently, and their benthic sources of food have not been investigated. A better understanding of mysid and crayfish benthic food preferences would allow for determination of their potential to impact special status communities in Lake Tahoe. In most other oligotrophic lakes, eutrophication has been known to increase benthic invertebrate densities through increased pelagic primary production and subsequent fallout to the benthos (e.g., Robertson and Alley 1966, Clarke et al. 1997, Nalepa et al. 2000). Comparisons of contemporary and historical collections suggest that the opposite has occurred in Lake Tahoe. Increased eutrophication may be causing increased shading to the benthos and associated declines in benthic plant abundance. It is possible that this decline in preferred native habitat for endemic invertebrates, along with an increase in Proposal: Tahoe Science Program Round 12 Request for Proposals numbers of non-native invertebrate taxa has resulted in the dramatic endemic invertebrate declines recently observed (Figure 5). The field collections and lab experiments proposed herein will allow for a greater understanding of Lake Tahoe’s unique special status communities and the mechanisms contributing to their declines. d. Goals, objectives, and hypotheses to be tested The goals of this proposal are: 1) increase our understanding of the biology and ecology of deepwater special status plant and invertebrate communities, 2) determine mechanisms (e.g. decreased water clarity and the introduction of non-native species) that have contributed to declines in these communities over the past 40+ years, and 3) to create a restoration and monitoring plan based on determined mechanisms. Specific objectives and related hypotheses include: Objective 1. Increase our understanding of special status aquatic plant (stoneworts, liverworts, and mosses) and invertebrate (Tahoe stonefly and blind amphipod) communities in deepwater “hotspot” areas of Lake Tahoe and relate the distribution of these communities to depth, subsurface irradiance, substrate type, and availability of organic matter in sediments. Hypothesis 1: The spatial extent of deepwater plant communities in Lake Tahoe is influenced by depth, substrate, and the amount of light reaching benthic habitats. Special status invertebrate distribution is determined by depth, substrate type (including plant presence or absence), and organic matter availability in sediments. Objective 2. Increase our understanding of the biology and ecology of special status deepwater plants and invertebrates through seasonal tracking of populations, developmental state, and diet using stable isotopes. Hypothesis 2. Deepwater stoneworts, mosses, and liverworts will be present throughout the year. The Tahoe stonefly and blind amphipod will have relatively long (≥1 year) life cycles, based on the deep, coldwater conditions of the targeted benthic habitats. Endemic invertebrates will be reliant on organic matter from pelagic sources when benthic sources of organic matter are not readily available. Objective 3. Develop a P-I curve for commonly encountered deepwater plants in Lake Tahoe and link the curve to historical light data to determine how plant communities have responded to changes in clarity and how they may respond to future changes in water clarity. Hypothesis 3. Plant distribution is restricted to a limited depth interval in Lake Tahoe that corresponds with subsurface irradiance levels sufficient for growth. Change in plant distribution over time is linked to the amount of light reaching benthic habitats and thus can be projected into the future according to load reduction scenarios in the Tahoe TMDL. Objective 4. Determine if non-native invertebrate species are affecting special status communities. Hypothesis 4. Introduced mysid shrimp and signal crayfish are both opportunistic crustacean omnivores in Lake Tahoe. The introduction and increase in abundance of these species in Lake Tahoe have affected deepwater special status communities through the alteration of habitat and direct predation. Objective 5. Identify a strategy to conserve and restore habitat for deepwater special status plant and invertebrate communities and develop a monitoring plan to evaluate community response to conservation and restoration strategies. Proposal: Tahoe Science Program Round 12 Request for Proposals Hypothesis 5. Improvements in water clarity and declines in non-native invertebrates will help restore special status communities in Lake Tahoe. e. Approach, methodology and location of research Objective 1. Deepwater plant and invertebrate communities will be sampled at two deepwater “hotspot” areas in Lake Tahoe. “Hotspot” areas are defined here as areas where endemic communities were located in recent surveys (McKinney Bay and South Lake Tahoe; Figure 6). Although deepwater plants and associated endemic stoneflies should be relatively easy to encounter in the targeted area in South Lake Tahoe, blind amphipods will likely be harder to find, given their poor representation in samples collected in 2008-09. At least five replicate samples will be collected with a Ponar grab at each depth for each location. Sampling effort will increase as the difficulty of encountering an organism increases. Initial surveys will occur in July and September to determine the spatial occurrence of benthic plants and invertebrates. If deepwater plants are not encountered with a Ponar grab, short benthic trawls will be conducted at discreet depths. At each location, measurements of subsurface irradiance will be taken with a Li-Cor underwater PAR sensor and Secchi depth will be recorded. Once plant and endemic distribution is determined for each location, areas with the highest densities of these taxa will be targeted for temporal analysis, as described in objective 2. Upon collection of a sample, live plant material will be removed and placed on ice for identification and morphological analysis in the laboratory. In the laboratory, plants will be identified, their general condition recorded, and selected characteristics such as length and photosynthetically active area will be measured. If justified by preliminary trials, we will also measure chlorophyll a and b and carotenoid concentrations (pigment per unit of plant biomass) because the absolute values as well their ratios are usually related to the light response of photosynthesis (Marschal and Proctor 2004). Plants will be dried to a constant biomass at 70° C and weighed. Once plants are removed, samples will be sieved through a 500 µm mesh bucket sieve. Material retained in the sieve will be picked for live invertebrates and invertebrates will be placed in 70% ethanol for laboratory analysis. All sediment will be kept, dried at 70° C to a constant biomass, sieved, and weighed to determine the relative percentage of each sediment category (silt, sand, gravel, and cobble) according to Wentworth (1922). Sediment will be combusted at 550° C for 2 h to determine the ash-free dry mass (AFDM) of organic material in each sample. Objective 2. Samples will be collected six times throughout the year (March, May, July, September, November, and January) from target locations where special status plant and invertebrates are found in initial surveys (objective 1). When encountered, endemic invertebrates will be measured (body length and head capsule width) and weighed in an attempt to determine seasonal size distribution for life history determinations. Endemic invertebrates will also be analyzed for stable isotopes to determine feeding habits (pelagic vs. benthic sources and trophic position). Plants will be identified, morphologically examined, dried and weighed as described in objective 1 to determine seasonal distribution and life history characteristics. Objective 3. Deepwater plants will be collected from areas where they are encountered during late summer to determine rates of photosynthesis rates under differing light conditions. Upon collection, a standard fresh weight of plant will be incubated in a BOD bottle filled with lake water for approximately two hours. For the duration of the experiment, bottles will be kept in a water bath equilibrated to the temperature of the bottom environment from which the plants were collected. Neutral density filters will be used to shade bottles, thus exposing plants to different irradiance levels corresponding to the measured light intensity along a vertical depth profile in Lake Tahoe. The triplicated treatments will include dark bottles to measure plant respiration and light bottles without plants as controls. Oxygen measurements will be taken with an oxygen electrode in the bottles before and after incubation to determine photosynthesis-irradiance (P-I) curves for the most commonly encountered deepwater plant taxa in Lake Tahoe (sensu Menendez and Sanchez 1998, Goldsborough and Kemp 1988). The relationship between Proposal: Tahoe Science Program Round 12 Request for Proposals the P-I curve obtained and long-term subsurface light data will allow for modeling of plant community responses to changing light conditions over time (past and future). In particular, we will use the predicted changes in lake transparency modeled from various load reduction scenarios in the Lake Tahoe TMDL to evaluate the impact on deepwater benthic plants. Objective 4. Non-native opossum shrimp (Mysis relicta) and signal crayfish (Pacifastacus leniusculus) will be collected 2-3 times per year in McKinney Bay and South Lake Tahoe. Opossum shrimp will be collected by dragging a large-diameter zooplankton net along the bottom of the lakeduring the day and placing collected individuals immediately formalin for gut content analysis. Crayfish will be collected with minnow traps and preserved in formalin immediately for gut content analysis. Additional opossum shrimp and crayfish will be collected and acclimated to laboratory conditions for use in laboratory experiments. In the laboratory, experiments testing opossum shrimp and crayfish predation on amphipods and stoneflies will be conducted. In each experiment, surrogates of endemic invertebrates will be used due to the difficulty and impact of collecting large quantities of endemic invertebrates. Common amphipods and stoneflies from other systems will be used to represent endemic Tahoe invertebrates. The size of surrogates will reflect juvenile and adult forms of each endemic taxon and caution will be taken in interpreting results given that surrogate prey items may differ slightly from actual special status species. One opossum shrimp will be placed in tanks with five potential prey items of the same taxon and experimental tanks will be checked at 1, 3, and 6 hours to determine prey consumption. Experimental treatments will be: no substrate, silt substrate, gravel substrate, and silt/plant substrate. Predation by crayfish on amphipods, stoneflies, and plants will be conducted using the same treatment types and laboratory conditions. Light and temperature conditions in the laboratory will be matched as closely as possible to field conditions. Objective 5. Information collected from field and laboratory analyses in objectives 1-4 will allow for determination of conservation and restoration strategies for deepwater special status communities and their habitat in Lake Tahoe. Mechanisms determined to have been most detrimental to deepwater special status communities in Lake Tahoe will be identified as target stressors that should be managed, if possible. A monitoring plan will be developed to allow managers to determine how deepwater special status communities might respond to restoration measures. The monitoring plan will include details about tracking these communities and their responses to changes in the benthic environment over time. f. Relationship of the research to previous and current relevant research, monitoring, and/or environmental improvement efforts Of critical importance is the direct connection between this proposed research and current efforts to develop and refine an environmental indicators program and a monitoring and evaluation program for Lake Tahoe. The Tahoe Regional Planning Agency and collaborating partners have had numerous meetings in the last seven years to (1) determine environmental condition, (2) assess response to restoration efforts, (3) monitor status and trends, and (4) establish scientifically-based targets for indicators of choice. The proposed research will contribute significantly to these emerging efforts by focusing on an area that has not been previously considered in Lake Tahoe. Previous funding (20092010) by the Tahoe Regional Planning Agency recognized our lack of understanding of the benthic environment and the importance of baseline community characterization given recent invasive species discoveries and eutrophication to the lake. This was the first major funding for benthic biodiversity studies since the 1960s. As previously noted, our scientific team has discovered dramatic changes to benthic assemblages at the lake bottom (see Figure 1), suggesting that we should pursue additional information about this deepwater environment, particularly where declines have occurred. We hope to refine our understanding of benthic processes to develop indicators that are reliable and sensitive to the Proposal: Tahoe Science Program Round 12 Request for Proposals changing conditions in Lake Tahoe. Other large lake ecosystems have undergone much greater alterations due to eutrophication (oligotrophic to eutrophic) and have documented subsequent changes to benthic plant and invertebrate communities. Surprisingly, Lake Tahoe has only undergone progressive eutrophication with a slight change in trophic status; thus restoration of habitat for deepwater special status communities is possible. g. Strategy for engaging with managers and obtaining permits We will engage managers by presenting quarterly updates to the funding agency and an annual presentation to agencies managing water quality thresholds within the basin (Tahoe Regional Planning Agency, Lahontan Water Quality Control Board, Nevada Division of Environmental Protection, California Department of Fish and Game, and the Nevada Department of Wildlife). We will also present at technical meetings, the Tahoe Science Symposium, and to the Science Management Integration team or other agency sponsored meetings as requested. We did not have to apply for permits for our previous whole lake survey project funded by the Tahoe Regional Planning Agency; thus, to our knowledge, we are not required to obtain permits. If this changes, we will comply with permitting needs and request additional funding to process these permits. Finally, we will create information fact sheets that can be placed on websites or handed out at the annual Tahoe summit in the summer time to inform policy makers and the public about the ecology and status of endemic species and their habitat. h. Description of deliverables/products and plan for how data and products will be reviewed and made available to end users A primary product of this project will be the determination of the mechanisms contributing to the decline of deepwater special status communities and development of recommendations to conserve and restore habitat for these communities. Once these mechanisms are determined, we will provide a monitoring plan that will allow for evaluation of special status community response to changes in variables associated with the declines (e.g. water clarity, density of invasive species). In the monitoring plan, we will provide specifics for future monitoring, such as optimal temporal and spatial targets for sampling and appropriate methodologies. Additionally, we will provide information related to the biology, status, and distribution of native deepwater aquatic plants and endemic invertebrates. A preliminary report (end of first year), and a final report (end of project) will be prepared. Throughout the project, we will be working in close cooperation with Shane Romsos, Acting Branch Chief of the Tahoe Regional Planning Agency, who has been working with other agencies to develop quantifiable indicators to address changes in the Lake Tahoe Basin. In order to facilitate the transfer of information to audiences beyond the Tahoe basin, we will present at annual scientific meetings (American Society of Limnology and Oceanography or the North American Benthological Society), and make public presentations of data as requested. Proposal: Tahoe Science Program Round 12 Request for Proposals III. Schedule of major milestones/deliverables Project Dates: June 1, 2012 – May 31 2014 Milestone/Deliverables Start Date End Date Prepare progress reports June 2012 April 2014 Description Submit brief progress report to Tahoe Science Program coordinator by the 1st of July, October, January, and April Prepare field equipment; Initiate field collections for dispersed deepwater benthic sampling Initiate field collections for focused seasonal sampling and determination of life history characteristics Process and identify plants and invertebrates from field collections. Process stable isotope samples Run P-I curve for common Lake Tahoe plants and process data Objective 1 Field Collections June 2012 October 2012 Objective 2 Field Collections October 2012 October 2013 September 2012 December 2013 Objective 3 July 2013 October 2013 Objective 4 October 2013 Objective 5 January 2014 January Run laboratory experiments with non-native 2014 invertebrates and process laboratory experiment data March Prepare restoration strategy recommendations 2014 and monitoring plan 1st annual accomplishment report September 2013 September Prepare annual summary of accomplishments 2013 Final presentation to basin managers April 2014 April 2014 Final presentation to basin managers seeking oral feedback for the final report Final report May 2014 May 2014 Final submission of our final report including an Executive Summary Objectives 1-2 Proposal: Tahoe Science Program Round 12 Request for Proposals IV. Literature cited/References Abrahamson, S.A.A., and C.R. Goldman. 1970. Distribution, density, and production of the crayfish Pacifastacus leniusculus Dana in Lake Tahoe, California-Nevada. Oikos 21:83-91. Bailey, R.J.E., J.T.A. Dick, R.W. Elwood, and C. MacNeil. 2006. Predatory interactions between the invasive amphipod Gammarus tigrinus and the native opossum shrimp Mysis relicta. Journal of the North American Benthological Society 25:393-405. Baumann, R.W. 1984. Review: aquatic insects. Ecology 67:589-590. Beauchamp, D.A., B.C. Allen, R.C. Richards, W.A. Wurtsbaugh, and C.R. Goldman. 1992. Lake trout spawning in Lake Tahoe: egg incubation in deepwater macrophyte beds. North American Journal of Fisheries Management 12:442-449. Caires, A., S. Chandra, M. Wittmann, and G. Schladow. 2010. Long-term change in benthic invertebrate assemblages in Lake Tahoe, California/Nevada. 5th biennial meeting abstracts-Lake Tahoe Basin Science Conference, Incline Village, Nevada. Chandra, S. 2011. Signal crayfish (Pacifastacus leniusculus) in Lake Tahoe: history, ecology, and potential management. Report to Nevada Board of Wildlife Commissioners. University of Nevada, Reno. Chandra, S., M.J. Vander Zanden, A.C. Heyvaert, B.C. Richards, B.C. Allen, and C.R. Goldman. 2005. The effects of cultural eutrophication on the coupling between pelagic primary producers and benthic consumers. Limnology and Oceanography 50:1368-1376. Clarke, K.D., R. Knoechel, and P.M. Ryan. 1997. Influence of trophic role and life-cycle duration on timing and magnitude of benthic macroinvertebrate response to whole-lake enrichment. Canadian Journal of Fisheries and Aquatic Sciences 54:89-95. Covich, A.P., and J.H. Thorp. 2001. Introduction to the subphylum Crustacea. Pages 777-809 in Ecology and classification of North American freshwater invertebrates (J.H. Thorp and A.P. Covich, editors). Second edition. Academic Press, San Diego, California. Flint, R.W. 1975. The natural history, ecology and production of the crayfish, Pacifastacus leniusculus, in a subalpine lacustrine environment. Ph.D. thesis, University of California, Davis. 150 pp. Frantz, T.C., and A.J. Cordone. 1966. A preliminary checklist of invertebrates collected from Lake Tahoe, 1961-1964. Biological society of Nevada occasional papers no. 8. 12 pp. Frantz, T.C., and A.J. Cordone. 1967. Observations on deepwater plants in Lake Tahoe, California and Nevada. Ecology 48:709-714. Frantz, T.C., and A.J. Cordone. 1996. Observations on the macrobenthos of Lake Tahoe, CaliforniaNevada. California Fish and Game 82:1-41. Goldman, C.R. 1988. Primary productivity, nutrients, and transparency during the early onset of eutrophication in ultra-oligotrophic Lake Tahoe, California-Nevada. Limnology and Oceanography 33:1321-1333. Goldman, C.R., M.D. Morgan, S.T. Threlkeld, and N. Angeli. 1979. A population dynamics analysis of the Cladoceran disappearance from Lake Tahoe, California-Nevada. Limnology and Oceanography 24:289-297. Goldsborough, J., and W.M. Kemp. 1988. Light responses of a submersed macrophyte: implications for survival in turbid tidal waters. Ecology 69: 1775-1786. Jassby, A.D., J.E. Reuter, R.P. Axler, C.R. Goldman, and S.H. Hackley. 1994. Atmospheric deposition of nitrogen and phosphorus in Lake Tahoe (California-Nevada). Water Resources Research 30:22072216. Jassby, A.D., J.E. Reuter, and C.R. Goldman. 2003. Determining long-term water quality in the presence of climate variability: Lake Tahoe (USA). Canadian Journal of Fisheries and Aquatic Sciences 60:1452-1461. Jewett, S.G., Jr. 1965. Four new stoneflies from California and Oregon (Plecoptera). Pan-Pacific Entomologist 41:5-9. Johannsson, O.E., M.F. Leggett, L.G. Rudstam, M.R. Servos, M.A. Mohammadian, G. Gal, R.M. Dermott, and R.H. Hesslein. 2001. Diet of Mysis relicta in Lake Ontario as revealed by stable isotope Proposal: Tahoe Science Program Round 12 Request for Proposals and gut content analysis. Canadian Journal of Fisheries and Aquatic Sciences 58:1975-1986. Linn, J.D., and T.C. Frantz. 1965. Introduction of the opossum shrimp (Mysis relicta Loven) into California and Nevada. California Fish and Game 51:48-51. Marschall, M., and M.C.F. Proctor. 2004. Are bryophytes shade plants? Photosynthetic light responses and proportions of chlorophyll a, chlorophyll b and total carotenoids. Annals of Botany 94:593–603. Menendez, M., and A. Sanchez. 1998. Seasonal variations in P-I responses of Chara hispida L. and Potamogeton pectinatus L. from stream Mediterranean ponds. Aquatic Botany 61:1-15. Nalepa, T.F., G.A. Lang, and D.L. Fanslow. 2000. Trends in benthic macroinvertebrate populations in southern Lake Michigan. Verhandlungen der Internationalen Vereinigung für Theoretische und Angewandte Limnologie. 27:2540-2545. Parker, J.I. 1980. Predation by Mysis relicta on Pontoporeia hoyi: a food chain link of potential importance in the Great Lakes. Journal of Great Lakes Research 6:164-166. Richards, R.C., C.R. Goldman, T.C. Frantz, and R. Wickwire. 1975. Where have all the Daphnia gone? The decline of a major cladoceran in Lake Tahoe, California-Nevada. Verhandlungen der Internationalen Vereinigung für Theoretische und Angewandte Limnologie. 19:835-842. Robertson, A., and W.P. Alley. 1966. A comparative study of Lake Michigan macrobenthos. Limnology and Oceanography 11:576-583. Rybock, J.T. 1978. Mysis relicta Loven in Lake Tahoe: vertical distribution and nocturnal predation. Ph.D. thesis, University of California, Davis. 116 pp. Sand-Jensen, K., and J. Borum. 1991. Interactions among phytoplankton, periphyton, and macrophytes in temperate freshwaters and estuaries. Aquatic Botany 41:137-175. Schwarz, A., Hawes, I., and Howard-Williams, C. 1996. The role of photosynthesis/light relationships in determining lower depth limits of characeae in South Island, New Zealand lakes. Freshwater Biology 35:69-80. Schwarz, A., Hawes, I., and Howard-Williams, C. 1999. Mechanisms underlying the decline and recovery of a characean community in fluctuating light in a large oligotrophic lake. Australian Journal of Botany 47:325-336. Schwarz, A., Howard-Williams, C., and J. Clayton. 2000. Analysis of relationships between maximum depth limits of aquatic plants and underwater light in 63 New Zealand lakes. New Zealand Journal of Marine and Freshwater Research 34: 157-174. Sealer, D.B., and F.P. Binowski. 1988. Vulnerability of early life intervals of Coregonus hoyi to predation by a freshwater mysid, Mysis relicta. Environmental Biology of Fishes 21:117-126. Swift, T.J., J. Perez-Losada, S.G. Schladow, J.E. Reuter, A.D. Jassby, and C.R. Goldman. 2006. Water clarity modeling in Lake Tahoe: Linking suspended matter characteristics to secchi depth. Aquatic Sciences 68: 1-15. TERC. 2011. Tahoe: 2011 State of the Lake Report. Tahoe Environmental Research Center, University of California, Davis. 79 pp. (terc.ucdavis.edu). Threlkeld, S.T. 1981. The recolonization of Lake Tahoe by Bosmina longirostris: evaluation the importance of reduced Mysis relicta populations. Limnology and Oceanography 26:433-444. Vander Zanden, M.J., S. Chandra, B.C. Allen, J.E. Reuter, and C.R. Goldman. 2003. Historical food web structure and restoration of native aquatic communities in the Lake Tahoe (California-Nevada) basin. Ecosystems 6:274-288. Wentworth, C.K. 1922. A scale of grade and class terms for clastic sediments. The Journal of Geology 30:377-392. Wilhelm, F.M., J. Hamann, and C.W. Burns. 2002. Mysid predation on amphipods and Daphnia in a shallow coastal lake: prey selection and effects of macrophytes. Canadian Journal of Fisheries and Aquatic Sciences 59:1401-1408. Zhu, B., D.G. Fitzgerald, C.M. Mayer, L.G. Rudstam, and E.L. Mills. 2006. Alteration of ecosystem function by zebra mussels in Oneida Lake: impacts on submerged macrophytes. Ecosystems 9:10171028. Proposal: Tahoe Science Program Round 12 Request for Proposals V. Figures Figure 1. A comparison of historical (1962-63) and contemporary (2008-09) data from benthic collections in Lake Tahoe. The comparisons show: a) lakewide mean±SE total invertebrate density, b) blind amphipod densities by depth, c) Tahoe stonefly densities by depth, and d) benthic plant occurrence at defined depth intervals. Proposal: Tahoe Science Program Round 12 Request for Proposals Figure 2. Mean±SE invertebrate density for each sediment category (dominant substrate) in benthic collections from the 1960s. Figure 3. Secchi depths recorded from 1968-2011 by UC Davis, showing the steady decline in water clarity since the late 1960s. Figure taken from TERC (2011). Proposal: Tahoe Science Program Round 12 Request for Proposals Crayfish (mean density/trap) 40 35 30 25 20 15 10 5 0 1967 1974 2008 Year Figure 4. Average lakewide density (total crayfish per trap) in Lake Tahoe for three time periods since 1967. Data presented are from Abrahamson and Goldman (1970), Flint (1975) and Umek et al. (unpublished data). Figure 5. A conceptual model showing potential interactions between deepwater special status species, their environment, and non-native species. One of the main goals of the proposed study is to determine which interactions are occurring how these interactions are affecting special status communities. Proposal: Tahoe Science Program Round 12 Request for Proposals Figure 6. A map of Lake Tahoe showing sampling locations from a 2008-09 benthic survey (black dots), sampling locations of a 1962-63 benthic survey (black arrows), and targeted locations for field surveys described in this proposal (outlined circular areas).