Proposal: Tahoe Research Supported by SNPLMA 2010 I. Title Page Title Subtheme Principal Investigator and Receiving Institution Co‐Principal Investigator Agency Collaborator Grants Contact Person Funding requested: Total cost share contributions): The ecology of curly leaf pondweed (Potamogeton crispus) and the potential for control using bottom barriers in Lake Tahoe 2a: Understanding the impacts of aquatic invasive species Marion Wittmann, Ph.D. University of California Davis Tahoe Environmental Research Center 291 Country Club Drive, Incline Village, NV 89451 Phone: 775 881 7560 x7402 Fax: 775 832‐1673 Email: mwittmann@ucdavis.edu Sudeep Chandra, Ph.D. Department of Natural Resources and Environmental Science Mail Stop 186 University of Nevada Reno, NV 89557 Phone: 775‐784‐6221 Fax: 775‐784‐4583 Email: sudeep@cabnr.unr.edu Lars W J Anderson USDA Agricultural Research Service Department of Plant Science, Mail Stop 4 Weed Science Program, UC Davis One Shields Avenue Davis, CA 95616 Phone: (530) 752‐7870 Fax: (530) 752‐4604 lars.anderson@ars.usda.gov Kim Boyd Tahoe Resource Conservation District 870 Emerald Bay Road, Ste. 108 South Lake Tahoe, CA 96150 Phone: 530 543 1501 x109 Fax: 530.543.1660 Email: kboyd@tahoercd.org Ted Thayer and Rita Whitney Tahoe Regional Planning Agency p.o. box 5310 Stateline, Nevada Phone: 775 589 5301 Fax: 775 588 4527 Email: tthayer@trpa.org George Malyj UC Davis John Muir Institute of the Environment (JMIE) Watershed Sci Bldg, RM 1105G, Davis, CA 95616 Phone: 530 752 3938, FAX: 530 754 9364 gjmalyj@ucdavis.edu $ 184,040 $ 1 Proposal: Tahoe Research Supported by SNPLMA 2010 II. Proposal Narrative a. Project abstract Aquatic invasive species introductions to Lake Tahoe have increased in recent decades and are rapidly dispersing and impacting the nearshore of Lake Tahoe. Curly leaf pondweed (Potamogeton crispus) was recently discovered in the southern portion of Lake Tahoe and is rapidly expanding along the littoral zone. Because of curly leaf pondweed’s recent introduction and restricted range, it is a viable candidate for control or eradication in Lake Tahoe. The understanding of the interaction between the ecology and management of an invasive species is key toward a successful control program. The major objectives of this proposal are to: (1) identify the role the “turion bank” of curly leaf pondweed plays in Lake Tahoe waters and the potential for this bank to contribute to the spread of the invasive species, (2) the susceptibility of this bank to the treatment of three kinds of bottom barriers, and (3) recommend and outline the method which should be employed at the lake to prevent further expansion of the plant. This proposal directly addresses the need for control and management of this new and aggressive invasive plant species. b. Justification statement: explain the relationship between the proposal and the subtheme(s) In order to control or eradicate Lake Tahoe’s established aquatic invasive species, it is necessary to understand the species‐specific ecology combined with novel applications of effective low cost non‐ chemical treatments to prevent population growth. Curly leaf pondweed (Potamogeton crispus), is one of Lake Tahoe’s most recent nearshore aquatic invaders, and because of its unique life history traits (turion production and seasonal growth patterns), this species has competitive advantages over native aquatic plants and has been called one of the most widely distributed, nuisance forming taxon in North America (Crowell 2003, Johnson 2007). Due to its recent introduction, and restricted range in the southern portion of Lake Tahoe (Figure 1), curly leaf pondweed is in an early invasion stage (Figure 2) and thus is a candidate for eradication or control. The goal of this research is to study a life history trait of curly leaf pondweed, specifically, vegetative turion production, its relationship to plant biomass, and the role that it plays in (1) population expansion in the nearshore of Lake Tahoe, (2) its susceptibility to non‐chemical treatment using gas permeable and impermeable bottom barriers, and (3) recommend the method(s) that should be employed to control this early invader that accounts for the potential rate of population growth in certain conditions in the lake. Through this research, we will provide information related to the invasion pressure of variable density populations of curly leaf pondweed in the lake and the contributions they have to range expansion in the nearshore. This will inform the potential for rapid propagation, and provide managers with a mechanism to identify the magnitude of an infestation as an invasion source and target specific densities of this species for control. In addition, we will determine the effectiveness of a currently utilized control mechanism (bottom barriers) on the reduction of curly leaf pondweed plant biomass as well as on turion germination. We will return to the experimental treatment sites one half year and one year after application and measure recolonization rates. Through this research, we can make recommendations regarding the use of a feasible technology for non‐chemical treatment of a new invasive species, recolonization rates associated with this treatment, and a life‐history based predictive model to assess the rate of population growth, areas to target for treatment based on differential population growth rates, and potential reductions to estimated growth rates based on management efforts. c. Concise background and problem statement Invasive species introductions and impacts in Lake Tahoe and the need for effective management. Aquatic invasive species have been identified in recent years as a major threat to Lake Tahoe and there have been a number of rapid responses by basin management agencies to monitor and control various non‐native invertebrate, macrophyte and warmwater fish populations. In particular, three recent 2 Proposal: Tahoe Research Supported by SNPLMA 2010 invaders, Asian clam (C. fluminea) (discovered 2002), Eurasian watermilfoil (Myriophyllum spicatum) (identified 1995, Anderson and Spencer 1996) and curly leaf pondweed (Potamogeton crispus) (discovered 2003) are aggressively spreading within Lake Tahoe and are establishing or threatening surrounding water bodies in the region. In Lake Tahoe, Asian clam has been shown to alter nutrient cycling and water quality by creating localized increases in nitrogen, phosphorus and calcium, and are also associated with filamentous algal blooms and decreases to nearshore aesthetic through shell matter deposition (Wittmann et al. 2008). In the Tahoe Keys, Eurasian watermilfoil provides habitat for warmwater invasive fishes (Kamerath et. al. 2008), pumps phosphorus into the water column potentially leading to increased algae production (Walter 2000), and is a navigation nuisance to homeowners, who incur an annual harvesting cost of $260,000 for this species (Dotson 2007). Similarly, curly leaf pondweed has spread across a majority of the south shore of Lake Tahoe since its discovery (Figure 1), and has been called the most widely dispersed, nuisance forming invasive aquatic macrophytes in North American lakes where it can inhibit recreation, increase water column phosphorus concentrations and impair water quality (Bolduan et al. 1994, Woolf and Madsen 2003). Bottom barriers as a non‐chemical management strategy for aquatic invasive species The use of benthic barriers to inhibit vital physiological or metabolic requirements such as photosynthesis or respiration is well known as a non‐chemical management strategy for aquatic nuisance species (Engel 1984, Ussery et al. 1997, Gunnison and Barko 1992, Eichler et al. 1995, Wittmann et al. submitted). In Lake Tahoe, the use of polyethylene fabric bottom barriers to control Eurasian watermilfoil has been implemented since the 2007 (Rita Whitney, TRPA pers. comm.) with localized population reductions after treatment. In 2009, UCD and UNR researchers, in cooperation with Tahoe basin agencies, field experimented with EPDM rubber benthic barriers on Asian clam populations and achieved 100% mortality in 28 days due to the reduction of dissolved oxygen concentrations under the barriers (Wittmann et al. submitted). In 2010, one acre of EPDM benthic barrier was applied to Asian clam populations in Lake Tahoe as part of a 120 day experiment using this method. Specific life history characteristics of Eurasian watermilfoil and Asian clam make these species ideal candidates for treatment with benthic barriers. Eurasian watermilfoil, as all submersed aquatic macrophytes, require light for growth and the use of benthic barriers manipulates the habitat by blocking light, resulting in both immediate and long term population reduction (USACE 2005). While Asian clam is tolerant to a number of toxicants, turbid conditions and wide temperature ranges, this species is hypoxia intolerant (Matthews and McMahon 1999) and thus populations can be reduced by eliminating dissolved oxygen with gas impermeable barriers. Traditional aquatic macrophyte (i.e., gas permeable) benthic barriers have been used on curly leaf pondweed in other systems with variable results. Mayhew and Runkel (1962) and Mayer (1978) reported that covering curly leaf pondweed populations with benthic barriers (polyethylene and Aquascreen™ (fiber glass coated sheeting)) was effective at reducing biomass, but did not report on subsequent recolonization. Madsen and Crowell (2002) suggest that bottom barriers are effective to prevent the growth of rooted curly leaf pondweed, however, long‐term management requires the elimination of vegetative buds called turions to interrupt the life cycle of curly leaf pondweed. The unique life history of curly leaf pondweed: Turion production, population propagation and the impacts of light and oxygen reduction Curly leaf pondweed sprouts from dormant shoot segments called turions. In the American northeast, turions are observed to germinate in late summer or fall, and the plants overwinter as small plants and resumes growth in spring when water temperatures rise (Figure 3). Individual stems may spread locally by rhizome growth and curly leaf pondweed biomass often reaches its maximum in early summer while flowering and fruiting occur from April to May. This seasonal pattern allows curly leaf pondweed to 3 Proposal: Tahoe Research Supported by SNPLMA 2010 avoid competition from other species, giving it ecological advantages as an invasive species in freshwater systems (Tobiessen and Snow 1984). Turion production is curly leaf pondweed main source of vegetative reproduction (Rogers and Breen 1980) and sexual reproduction occurs through seed production (Waisel 1971). However, seed germination is extremely low (<0.1%) and not commonly thought of as an important source of recruitment (Rogers and Breen 1980). Additionally, turions also act as a storage unit for carbohydrate (Madsen and Crowell 2002), which can enable dormancy until environmental conditions are more favorable. Though curly leaf pondweed is known to be susceptible to contact herbicide treatment, plant biomass regrowth often occurs due to turion formation as the major form of re‐infestation during the following growing season (Netherland et al. 2000). Thus, curly leaf pondweed turions are not only the main means of population propagation, reproduction and dispersal, but also can expand the window of opportunity for successful recruitment for this species through carbohydrate storage, herbicide evasion and season based competition. While turion germination can enable the sustainability of curly leaf pondweed populations, it can be limited by a number of physical factors. Laboratory and field studies have found that germination is generally controlled by temperature, light intensity and photoperiod (Rogers and Breen 1980, Sastroutomo 1981, Kadono 1982, Tobiessen and Snow 1984, Jian et al. 2003). However, in Asian systems where curly leaf pondweed is native and used to restore degraded lakes, turion germination has ceased as a result of low light availability and anoxic conditions (Wu et al. 2009, Short et al. 1987, Lauridsen et al. 1993, Clarke and Wharton 2001, Ni 2001). Understanding turion germination behavior is key to assessing the population dynamics of this species and the potential for non‐chemical treatment in Lake Tahoe. d. Goals, objectives, and hypotheses to be tested The goals of this study are to employ field and laboratory methods to understand the impact of curly leaf pondweed on Lake Tahoe’s littoral zone ecology and the potential for non chemical treatment using benthic barriers. Specifically we would like to understand the how the life history of curly leaf pondweed impacts (a) the likelihood of population growth in a large, subalpine oligotrophic lake given its current distribution, biomass and turion bank, and (b) its ability to be effectively managed in Lake Tahoe with benthic barriers to reduce both macrophyte biomass as well as turion viability. We seek to achieve these goals by pursuing the following objectives: 1. To assess the potential for curly leaf pondweed population growth and risk of further establishment by quantifying: plant biomass and stem density in multiple locations in south Lake Tahoe, the asexual reproductive potential of curly leaf pondweed by quantifying the turion bank as it relates to plant biomass/density, and habitat quality. 2. We will use a compliment of laboratory and field studies to assess the impact of three different bottom barrier fabrics on curly leaf pondweed vegetation and turion germination. Sediment anoxia and light reduction has been shown to inhibit turion germination of curly leaf pondweed in its native range (Wu et al. 2009). We will apply two gas permeable fabrics (polyethylene and jute) and one gas impermeable fabric (EPDM) to test the relationship between anoxia, light reduction and turion germination. 3. To evaluate the recolonization rate of curly leaf pondweed 6 months and 1 year after treatment with variable bottom barrier fabric types. 4. To predict the population recruitment rate of curly leaf pondweed based on plant biomass, habitat quality, and turion germination in Lake Tahoe under natural and managed (i.e., bottom barriers) conditions in order to make recommendations for non‐chemical or chemical control of this species. 4 Proposal: Tahoe Research Supported by SNPLMA 2010 5. Indentify specific regions in Lake Tahoe that present the greatest curly leaf pondweed invasion pressure based on life history traits and make recommendations for areas and methods of non‐ chemical treatments. The specific hypotheses to be tested are as follows: 1. Curly leaf pondweed plant biomass is dependent on habitat variability (nitrogen, phosphorus, dissolved inorganic carbon, pH, turbidity, temperature, light). 2. Curly leaf pondweed turion density is dependent on plant biomass. 3. Bottom barriers that reduce light permeability will reduce curly leaf pondweed macrophyte biomass and not turion germination rates. 4. Bottom barriers that reduce light permeability and dissolved oxygen concentrations will reduce curly leaf pondweed macrophyte biomass and turion germination. 5. Bottom barriers that reduce light permeability and dissolved oxygen concentrations will have a lower rate of recolonization than barriers that only reduce light permeability. 6. Based on the turion bank and habitat quality, curly leaf pondweed populations are expected to increase in Lake Tahoe. 7. Effective treatment that targets both biomass and turion viability reduction will reduce overall curly leaf pondweed population range in Lake Tahoe. e. Approach, methodology and location of research Approach: We will utilize a combination of field and laboratory surveys and experiments to understand the relationship between curly leaf pondweed population biomass, asexual reproduction (via turion production), and management using bottom barriers. This research will utilize a field survey to assess curly leaf pondweed plant and turion density in five locations in South Lake Tahoe, measure habitat quality in each location, field installation of three types of barriers to test impacts on plant biomass and turion viability, and the collection of turions from both control sites and from under barriers to quantify germination rates in a laboratory setting. Methodology and location: To acquire quantitative estimates for curly leaf pondweed vegetative biomass, we will use SCUBA to place 1 m2 quadrats (N = 5) to collect stem counts in five locations with variable density of curly leaf pondweed populations: (Ski Run marina, Tahoe Keys marina, Lakeside marina, outside of Tahoe Keys marina (open lake population), outside of Ski Run marina (open lake population). We will seek monoculture stands of curly leaf pondweed, however, due to the presence of other submersed macrophyte species in these areas, we will note all species present in each grid and their relative abundance. Following methods of Eichler et al. (1995), a Daubenmire scale (1968) will used to determine abundance classes in addition to quantitative estimates of density. Shoot biomass will be collected from each quadrat, dried (55°C), weighed and analyzed for tissue nutrient concentrations (carbon, nitrogen, and phosphorus). To characterize habitat at each location sediment particle size distribution, DIC, pH, turbidity, and sediment porewater and water column samples will be collected to measure nitrogen and phosphorus concentrations. Two temperature probes (iButtons®) will be installed at each location to measure temperature for a 1 year period. These values will be used as independent variables to predict curly leaf pondweed biomass and density (dependent variables) to test for habitat driven spatial differences in populations. Using methods of Johnson (2007) we will estimate the turion bank in Lake Tahoe. A petite Ponar sampler device (Wildco, Inc.) will be used to carry out the sediment turion survey in autumn 2011 in order to target the period of highest turion abundance and least plant 5 Proposal: Tahoe Research Supported by SNPLMA 2010 biomass (for ease of sampling) prior to ice formation (in marina sites). Three types of bottom barrier fabric will be installed at three curly leaf pondweed locations: polyethylene fabric (already used in Lake Tahoe for Eurasian watermilfoil treatment, blocks light but is gas permeable), EPDM rubber barrier (used in Lake Tahoe for Asian clam treatment, will block light and gas impermeable), and jute (a biodegradable material that will block a majority of light, however, increases BOD to decrease DO concentrations). Fifteen turions (collected from the immediate area) will be placed underneath each barrier to ensure their presence and availability for laboratory testing after barriers are removed. An in situ dissolved oxygen probe (Zebra Tech D‐Opto Logger, Accuracy: 1% or 0.02 PPM) will be placed under each barrier type and also in the control condition. Plant biomass and density estimates as well as barrier installation will occur in July 2011 (abundant curly leaf pondweed biomass present at this time in Lake Tahoe, due to unique seasonality of this species in this region, Lars Anderson pers. comm.) All barriers will be removed in September 2011 and estimates of any plant biomass and density underneath each barrier site plus one control site (untreated) will be taken. We will attempt to recover a total of 180 turions: 15 from an adjacent control plot (N = 1), as well as the 15 placed underneath each barrier type (N = 3), at each location (N = 3). The collected turions will be assessed for dormant or non‐dormant status (Sastroutomo 1981) and returned to the laboratory for viability analysis. Following methods of Sastroutomo (1980, 1981) we will estimate germination rates of turions from each barrier type warm treatment (35°C) in dark for 4 weeks before germination at 20°C and 12 h light (Bouldan et al. 1994). We will return to barrier treatment plots 6 months and 1 year after removal to observe recolonization rates of the variable barrier types. To model population growth and expansion of curly leaf pondweed in Lake Tahoe, we will use biomass and density estimates, habitat quality measurements, annual water temperature, and germination rates of treated and untreated turions to utilize a recruitment‐based population viability analysis (Morris and Doak 2003) to estimate the 5, 10 and 15 year estimates of population size for curly leaf pondweed. This population model will have two scenarios: one in which (benthic barrier) management occurs (impacts to plant biomass reduction and/or turion viability dependant on laboratory experimentation) and one in which the population is allowed to increase without control method is applied. f. Relationship of the research to previous and current relevant research, monitoring, and/or environmental improvement efforts Dr. Marion Wittmann (PI) is currently a project partner with the Tahoe Resource Conservation District, Tahoe Regional Planning Agency, and California State Parks in the Emerald Bay Eurasian watermilfoil control program. She and Dr. Sudeep Chandra (co‐PI) are also members of the Nearshore Aquatic Weed Working Group. The proposed research builds upon an ongoing program in Lake Tahoe to control Eurasian watermilfoil using a combination of bottom barriers and hand removal. However, this is new and novel research since no direct scientific investigations have occurred on this invasive species to date in Lake Tahoe. By observing the densities of the curly leaf pondweed turion bank as it relates to plant biomass in the field, a predictive model of population expansion can be used to design a management strategy (e.g., site selection based on source versus satellite populations) for this species. Additionally, while it is known that gas permeable barriers can reduce plant biomass in other systems, it has not shown to be an effective long term treatment for this species due to turion based recruitment. As discussed above, the reduction of light availability and dissolved oxygen concentrations has shown to inhibit turion germination, and the use of gas impermeable barriers to potentially control this species could be a low‐cost resource to managers in the basin in the current chemical‐free AIS treatment program. This methodology could ultimately improve the ecological health and aesthetic of the nearshore zone. This proposal is directly related to the conservation element of the TRPA regional plan 6 Proposal: Tahoe Research Supported by SNPLMA 2010 update. Specifically, as cited in the wildlife and fisheries sub‐element, this proposal lends directly towards the continued work with state and federal managers in the response to aquatic invasive species through the study of implemented management actions. Additionally, there is a nearshore component to the TRPA regional plan update. While the details of this component have not been supplied to the public by the TRPA, invasive macrophyte populations of Lake Tahoe are already having an impact to the nearshore water clarity and navigability via biomass accumulation. The efficient management and reduction of invasive aquatic macrophyte populations has the potential to remediate these impacts to the Tahoe nearshore. g. Strategy for engaging with managers and obtaining permits We will engage with managers by presenting results to the Lake Tahoe Aquatic Invasive Species Coordination Committee (LTAISCC) on a biannual basis, with a presentation of final findings at the project conclusion. We will attend Nearshore Aquatic Weed Working Group (NAWWG) meetings to present findings and provide information related to curly leaf pondweed location and management in Lake Tahoe. We will present findings to the Lake Tahoe Science Symposium in 2012. We will continue to work closely with Lars Anderson of the USDA Agricultural Research Service Exotic and Invasive Weed Research to ensure that this experimentation compliments ongoing USDA‐ARS curly leaf pondweed experimentation. This project will require permits. We will work closely with Kim Boyd of Tahoe Resource Conservation District and Ted Thayer and Rita Whitney of the Tahoe Regional Planning Agency to obtain the necessary field based permits for barrier application from the Lahontan Regional Water Quality Control Board, the Army Corps of Engineers and the California Department of State Lands. All proposed experimental sites are in California and therefore no Nevadan agencies will need to issue a permit for this work. Annual permits for Eurasian watermilfoil treatment with benthic barriers are already held by the Tahoe Resource Conservation District, and at the time of this proposed project period, those permits will be amended to incorporate regions of the Lake specific to this project. 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 research will be the understanding of the use of gas impermeable barriers for use as an eradication (rather than population reduction) tool for the management of curly leaf pondweed in Lake Tahoe. Low light availability and reduced dissolved oxygen concentrations in other systems has shown to inhibit turion germination; the ability to artificially create these conditions in Lake Tahoe can provide a feasible method for the removal of this early invasion stage nuisance plant. We will provide the results as a report showing the effectiveness of variable barrier fabrics, as well as the costs associated with implementing this strategy. A secondary product of this research will be an identification of the population growth potential of curly leaf pondweed in Lake Tahoe based on a unique life history trait, the vegetative turion. Once the population growth model is developed, we will provide estimates of population density based on habitat quality and management effort. We will be able to create a long‐ term (multi‐year) scenario of population change that can be used to identify priority areas for management based on life history traits. We will be working closely with Kim Boyd (TRCD) and Ted Thayer and Rita Whitney (TRPA) who have been working on the development of a comprehensive management strategy for invasive aquatic macrophytes in the basin. A preliminary report (end of year 1) and a final report (end of year 2) will be prepared and findings presented to members of the Lake Tahoe Nearshore Aquatic Weed Working Group. Finally, we will publish these results in a peer reviewed journal and present findings to academic and management motivated meetings (Western Aquatic Plant Management Society and National Lake Management Society). 7 Proposal: Tahoe Research Supported by SNPLMA 2010 III. Schedule of major milestones/deliverables Milestone/Deliverables Start Date End Date Initiate June 1, 2011 June 15, 2011 project/establish field plan Prepare quarterly July 2011 April 2013 progress reports Initiate field collections: plant survey and barrier installation/removal Finalize field collections, begin laboratory experimentation: turion germination Annual accomplishment report Description Plan field surveys and communication with Forest Service Submit brief progress report to Tahoe Science Program coordinator by the July 1, 2011, October 1, 2011, January 1, 2012, and April 1, 2012, July 1, 2011 June 2011 November 2011 Objectives 1, 2: Field characterization of pondweed populations, biomass and density estimates habitat quality sample collection, installation and removal of barriers November March 2012 Objective 2: Germinate turions from field/barrier 2011 collections to test for viability or germination rates as a result of barrier treatment, Field collections using Ponar to quantify turion bank counts September 2011 September Prepare annual summary of accomplishments in 2012 September. Recolonization observations May 2012 November 2012 Objective 3: Six month and 12 month observation of recolonization in barrier plots Data synthesis and predictive model building December 2012 Final reporting March 2013 March 2013 Objectives 4, 5: Begin data summarization and analysis, build population viability model based on field and laboratory collections June 2013 Draft final report (to be submitted in May 2013) and Final Report (to be submitted June 2013) 8 Proposal: Tahoe Research Supported by SNPLMA 2010 IV. Literature cited/References Anderson, L.W.J., Spencer, D. 1996. Survey of Lake Tahoe for presence of Eurasian watermilfoil. USDA Agricultural Research Service Aquatic Weed Control Investigations, Annual Report. Dept. of Vegetable Crops, Weed Science Program, UC Davis. Baker, H.G. 1989. Some aspects of the natural history of seed banks. In: Leck, M.A., V.T. Parker, and R.L. Simpson eds. Ecology of Soil Seed Banks. Academic Press, Inc. San Diego, USA. pp 9‐21. Boulduan, B.R., G.C. Van Eeckhout, H.W. Quade and J.E. Gannon. 1994. Potamogeton crispus the other invader. Lake and Reservoir Management 10:113‐125. Clark, S.J., Wharton, G. 2001. Sediment nutrient characteristics and aquatic macrophytes in lowland English Rivers. Science of the Total Environment 266:103‐112. Crowell, W. 2003. Curlyleaf pondweed: New Management Ideas for an Old Problem. Notes to the Minnesota Department of Natural Resources Exotic Species Program. http://www.lakewashingtonassn.com/pdfs/curlyleaf.pdf Daubenmire, R. 1968. Plant Communities: A Textbook of Synecology. Harper and Row, New York. 300 pp. Dotson, H. 2007. Aquatic weed management in Tahoe Keys. Presentation at the Western Aquatic Plant Management Society, Tahoe City, CA. Eichler LW, Bombard RT, Sutherland JW, Boylen CW (1995) Recolonization of the littoral zone by macrophytes following the removal of benthic barrier material. Journal of Aquatic Plant Management 33: 51‐54 Engel, S. 1984. Evaluating stationary blankets and removable screens for macrophyte control in lakes. Journal of Aquatic Plant Management 22: 43‐48. Gunnison D, Barko JW (1992) Factors influencing gas evolution beneath a benthic barrier. Journal of Aquatic Plant Management 30: 23‐28. Hobbs RJ, Humphries SE. 1995. An integrated approach to the ecology and management of plant invasions. Conservation Biology 9(4): 761‐770. Jian, Y.X., B. Li and J.B. Wang. 2003. Control of turion germination in Potamogeton crispus. Aquatic Botany 75:59‐ 69. Johnson, JA. 2007. Curlyleaf pondweed turion survey report, Roberds Lake. Report prepared for the Cannon River Watershed Partnership by Freshwater Scientific Services, LLC. Kadono, Y., 1982. Germination of the turion of Potamogeton crispus L. Physiology and Ecology of Japan 19: 1‐5. Kamerath, M., Chandra, S., Allen, BC. 2008. Distribution and impacts of warm water fish in Lake Tahoe USA. Aquatic Invasions (2008) Volume 3, Issue 1: 35‐41. Lauridsen, T.L., Jeppesen, Andersen, F.O. 1993. Colonization of submerged macrophytes in shallow fish manipulated Lake Vaeng: impact of sediment composition and waterfowl grazing. Aquatic Botany 46: 1‐15. Madsen, J.D., Crowell, W. 2002. Curleaf pondweed (Potamogeton crispus L.). Lakeline. Spring 2002. Pp 31‐32. Matthews M.A., McMahon R.F. 1999. Effects of temperature and temperature acclimation on survival of zebra mussels (Dreissena polymorpha) and Asian clams (Corbicula fluminea) under extreme hypoxia. J. Moll. Stud. 65: 317–325. Mayer, J. R., 1978. Aquatic Weed Management by Benthic Semibarriers. J. Aquat. Plant Manage. 16:31‐33. Mayhew, J.K., Runkel, S.T. 1962. The control of nuisance aquatic vegetation with black polyethylene plastic. Proc. Iowa Acad. Sci. 69: 302‐307. Morris, W. F., Doak, D. F. 2003. Quantitative Conservation Biology: Theory and Practice of Population Viability Analysis. Sinauer Associates, Sunderland, Massachusetts, USA Netherland, M.D., J.D. Skogerboe, C.S. Owens, and J.D. Madsen. 2000. Influence of water temperature on the efficacy of diquat and endothall versus curlyleaf pondweed. J. of Aquatic Plant Management. 38:25‐32. Ni, L. 2001. Stress of fertile sediment on the growth of submersed macrophytes in eutrophic waters. Acta Hydrobiologia Sinica 25: 399‐405. Rogers, K.H. ,Breen, C.M. 1980. Growth and reproduction of Potamogeton crispus in a South African Lake. Journal of Ecology. 68(2):561‐571. Sastroutomo, S.S. 1980. Environmental control of turion formation in curly pondweed (Potamogeton crispus). Physiol. Plant. 49:261‐264. Sastroutomo, S.S. 1981. Turion formation, dormancy and germination of curly pondweed, Potamogeton crispus L. Aquatic Botany 10: 161‐173. 9 Proposal: Tahoe Research Supported by SNPLMA 2010 Short, F.T. 1987. Effects of sediment nutrients on seagrasses: literature review and mesocosm experiment. Aquatic Botany 27:41‐57. Tobiessen, P. , Snow, P. D. 1984. Temperature and light effects on the growth of Potamogeton crispus in Collins Lake, New York State. Can. J. Bot. 62:2822‐2826. USACE 2005. U.S. Army Corps of Engineers Engineer Research and Development Center. Aquatic Plant Information System (APIS). http://el.erdc.usace.army.mil/aqua/apis. Accessed October 2010. Ussery, T.A., Eakin, H.L., Payne, B.S., Miller, A.C., Barko, A.W. 1997. Effects of Benthic Barriers on Aquatic Habitat Conditions and Macroinvertebrate Communities. J. Aquat. Plant Manage. 35: 69‐73. Waisel, Y. 1971. Seasonal activity and reproductive behavior of some submerged hydrophytes in Israel. Hydrobiologia. 12:219‐227. Walter, K. M. 2000. Ecosystem Effects of the Invasion of Eurasian Watermilfoil (Myriophyllum spicatum) at Lake Tahoe, CA‐NV. M.S. thesis, University of California, Davis Wittmann, M.E., Reuter, J.E., Chandra, S., Schladow, S.G., Allen, B.C., Caires, A. 2008. Asian clam (Corbicula fluminea) of Lake Tahoe: Preliminary Scientific Findings in Support of a Management Plan, Wittmann, M.E., Reuter, J.E., Chandra, S., Schladow, S.G., Allen, B.C. 2010. The control of an invasive bivalve Asian clam (Corbicula fluminea) using benthic barriers in a large natural lake. Submitted to Biological Invasions. Woolf, T.E., Madsen, J.D. 2003. Seasonal biomass and carbohydrate allocation patterns in Southern Minnesota curlyleaf pondweed populations. Journal of Aquatic Plant Management. 41: 113‐118. Wu, J., Cheng, S., Liang, W., He, F.,Wu, Z. 2009. Effects of sediment anoxia and light on turion germination and early growth of Potamogeton crispus. Hydrobiologia 628:111‐119. 10 Proposal: Tahoe Research Supported by SNPLMA 2010 V. Figures Figure 1. The distribution of Eurasian watermilfoil (Myriophyllum spicatum) and curly leaf pondweed (Potamogeton crispus) in Lake Tahoe, 1995 – 2006. Red circles indicate the presence of M. spicatum, yellow triangles indicate the presence of curly leaf pondweed, which was discovered in 2003. Map created and provided by Lars Anderson, USDA‐ARS. 11 Proposal: Tahoe Research Supported by SNPLMA 2010 Figure 2. A schematic of the theoretical relationship between the abundance of an invasive species over time in relation to management stage (Hobbs and Humphries 1995). When population numbers are low early in a species invasion, the rate of invasive species abundance changes slowly because of limits to reproductive capacity per unit time. These low abundance periods occur during an invasion where quarantine, eradication and control are priorities for managers because of the potential for management when invasive species abundance is low. Once populations have increased over an extended amount of time, effective control is unlikely without massive resource inputs. 12 Proposal: Tahoe Research Supported by SNPLMA 2010 Figure 3. Curly leaf pondweed life cycle as observed in North America (Madsen and Crowell 2002). Seasonal patterns observed in Lake Tahoe have different temporal scales than plant populations observed in the northeastern regions of North America (Lars Anderson, pers comm.) 13