DOES CLIMATE TRUMP FISH OR FISH TRUMP CLIMATE? LONG-TERM ECOLOGICAL DYNAMICS OF SUBALPINE CASTLE LAKE Sudeep Chandra, Wendy Trowbridge, Rene Henery, and Charles R. Goldman Outline • Lakes as sentinels of climate change – Physical and biological alterations linked to local and regional processes • Castle Station history: now the America’s oldest, continuous, mountain lake, ecological dataset • Long-term dynamics of climate, environmental, and biological variables • What is the influence of climate and fish manipulations on the zooplankton community composition? Federal Institute of Aquatic Science and Technology (Eawag), Dübendorf, Switzerland hLaboratory of Aquatic Photobiology and Plankton Ecology, Institute of Ecology, University of Innsbruck, Innsbruck, Austria iLimnological Institute, University of Konstanz, Konstanz, Germany jDepartment Lakes as sentinels of climate of Aquatic Food Webs, Netherlandschange Institute of Ecology, Centre for Limnology, k Nieuwersluis, The Netherlands Department of Aquatic Sciences and Assessment, Swedish University of Agricultural Sciences, and Department Ecology and Evolution,B.Uppsala a,*, Catherine b, HoracioofZagarese c, Stephen Rita Adrian M. O’Reilly Baines d, Dag O. l University, John Muir Institute of the Tahoe Environmental g,Environment, h, Dietmar Straile i, Hessene,Uppsala, Wendel Sweden Kellerf, David M. Livingstone Ruben Sommaruga Research Center, jUniversity of California, Davis, k California l Ellen Van Donk , Gesa A. Weyhenmeyer , and Monika Winder Abstract aLeibniz-Institute b of Freshwater Inland Fisheries, Berlin, Germany Biology While there is a general sense thatEcology lakesc canand act as sentinels of climate change, their efficacy has Program, Bard Annandale, New Laboratorio de Ecología y Fotobiología Acuática, notCollege, been thoroughly analyzed. WeYork identified the key response variables within a lake that act as Instituto Tecnológico (INTECH), Chascomús Provincia de Buenos Aires, Argentina indicators ofdetheChascomús effects of climate change on both the lake and the catchment. These variables dDepartment of Ecology and Evolution, Stony Brook University, Stony Brook, New York reflect a wide range of physical, chemical, and biological responses to climate. However, the eDepartment of Biology, University of Oslo, Oslo, Norway fCooperative Freshwater Ecology Unit, efficacy of the different indicators is affected by regional response to climate change, Ontario Ministryof ofthe thecatchment, Environment, Laurentian University, Sudbury,indicators Ontario,orCanada gSwiss characteristics and lake mixing regimes. Thus, particular Federal Instituteofof Aquaticare Science and Technology Switzerland combinations indicators more effective for different(Eawag), lake types Dübendorf, and geographic regions. The hLaboratory extraction of can be further by the influence of other environmental of climate Aquaticsignals Photobiology andcomplicated Plankton Ecology, Institute of Ecology, University of changes, such as eutrophication or acidification,Institute, and the equivalent reverse phenomena, in Innsbruck, Innsbruck, Austria iLimnological University of Konstanz, Konstanz, Germany jDepartment addition to of other land-use influences. many cases, however, canfor beLimnology, Aquatic Food Webs,InNetherlands Institute confounding of Ecology,factors Centre addressed through analytical toolskDepartment such as detrending or filtering. Lakes and are effective sentinelsSwedish for Nieuwersluis, The Netherlands of Aquatic Sciences Assessment, climate change because they are sensitive climate, respond rapidly to change, and integrate University of Agricultural Sciences, and to Department of Ecology and Evolution, Uppsala l information about changes in the catchment. University, Uppsala, Sweden John Muir Institute of the Environment, Tahoe Environmental Research Center, University of California, Davis, California Currently, climate change is considered to be one of the most severe threats to ecosystems Abstract around the globe (ACIA 2004; Rosenzweig et al. 2007 ). Monitoring and understanding the effects of climate change pose challenges because of the multitude of responses within an While there is a general sense thatvariation lakes canwithin act asthe sentinels of climate change, their ecosystem and the spatial landscape. A substantial body of efficacy research has not been demonstrates thoroughly analyzed. We identified key response variables withinchemical, a lake that the sensitivity of lakes tothe climate and shows that physical, andact as indicatorsbiological of the effects of climaterespond changerapidly on bothtothe lake and thechanges catchment. These variables lake properties climate-related ( ACIA 2004; (Limnology and Oceanography 2009) Regional climate influences by ocean cycles influences physical structure, heat content of lakes “All of the El Niño years and several others shows that the depth of the mixed layer and the mixing of heat into the stratified thermocline region control the storage of heat.” The storage of heat controls the production in a lake. (Strub et al., Science, 1985) Lake Washington case study: influence of thermal structure on primary production and resulting changes to zooplankton Changes in the thermal stratification Shifts in the onset of diatom blooms Select zooplankton species were influenced by alterations to the thermal regime Keratella- no Daphnia- yes (Winder and Schindler 2004) Outline • Lakes as sentinels of climate change – Physical and biological alterations linked to local and regional processes • Castle Station history: now the America’s oldest, continuous, mountain lake, ecological dataset • Long-term dynamics of climate, environmental, and biological variables • What is the influence of climate and fish manipulations on the zooplankton community composition? Castle Lake- one of 11 lakes in the Upper Sacramento region • • • • • • subalpine cirque lake mesotrophic maximum depth- 35 m ~ 54 % littoral habitat 2-4 year HRT Fish stocking across the Western mountains if they survived at Castle. Castle Lake, small subalpine lake located in the Klamath mountain range in California A test lake by agencies to determine what hatchery raised species would survive • • • • • • • • • • • Atlantic salmon Pacific salmon Arctic grayling Brown trout Rainbow trout Lake trout Brook trout Bass Catfish Bluegill Yellowstone cutthroat trout A young limnologist, arriving from Alaska, Dr. Charles Goldman recommends an investigation of the factors contributing to lake production, the program was initiated in 1958 and continues today Castle Lake measurements Climate Snow ice water content Snow depth Air temperatures Solar radiation Limnological factors (0-35 m) Temperature Dissolved Oxygen Vertical extinction coefficient Nitrogen Phosphorus Nutrient limitation assays Pelagic PPr- 14C Chlorophyll a Zooplankton composition/ biomass Fish stocking records ≈ 15 times in Summer, 1 Fall, 1 Winter/ Spring condition Conditions at Castle Lake can vary greatly depending on the season. ice free season, June- November 4000 Current fishes in the lake Biomass of Catchable fish stocked No trend 3500 3000 2500 2000 1500 1000 500 0 600 1959 500 1969 1979 1989 1999 2009 1979 1989 1999 2009 Biomass of Fingerling fish stocked p = 0.28, 16 pound decrease over the 52 years 400 300 200 100 0 1959 1969 • Brook trout (Salvelinus fontinalis)- naturalized spawner • Rainbow trout (Oncorhynchus mykiss)largely maintained through stocking with some limited natural recruitment • Golden shiner (Notemigonus crysoleuca)- forage fish with natural recruitment Late summer pelagic PPr in the mixed layer is influenced by zooplankton grazing pressure governed by trout, deep PPr variability results from changes in climate condition (Jassby et al. 1990, L&O) Current working conceptual model for understanding linkages between climate, fisheries stocking, and carbon transfer across the landscape Outline • Lakes as sentinels of climate change – Physical and biological alterations linked to local and regional processes • Castle Station history: now the America’s oldest, continuous, mountain lake, ecological dataset • Long-term dynamics of climate, environmental, and biological variables • What is the influence of climate and fish manipulations on the zooplankton community composition? Objectives and methods for this analysis 1) Evaluate the interactions between two human driven influences, climate and fish manipulations 2) How important is climate and lake phenological characteristics in understanding lake primary production? 3) How much does climate explain a shift in planktonic, primary consumers (zooplankton) during a period of experimental fish manipulation and drought period? 3) What drives zooplankton community structure before and after this period and do lakes recovery from these perturbations? Long-term trend analysis using regression corrected for autocorrelation, some of the environmental data is inherently nonlinear Created a hypothesized model to determine the influence of climate and fish manipulation on zooplankton structure. Divided data into 3 periods and used structural equation modeling (SEM), multivariate approach to look at a network of variables including latent variable, to understand the drivers of planktonic biomass. 1958-1987 Pre major fish manipulation, climate driven period 1987-1994 “Experimental” fish manipulation and extended drought conditions 1995-2011 Climate driven manipulation with intermittent fish stocking and non stocking periods 4 1 0 1959 -1 1969 1979 1989 1999 2009 -2 -3 -4 -5 180 140 120 80 60 40 20 140 1979 1989 1999 Snow Water Content p = 0.54, 8.1 cm increase over the 52 years 100 80 60 40 20 0 1959 1969 1979 1989 1999 140 120 100 80 1959 2009 1969 1979 1989 1999 2009 Minimum air temperature in August p = 0.0002, 4 degree increase over the 52 13 12 11 10 9 8 7 6 5 4 1959 2009 Winter condition 120 1969 160 34 1969 1979 1989 1999 2009 Maximum Air Temperature in August p = 0.02, 2.2 degree increase over 52 years 32 30 28 26 24 22 20 1959 1969 1979 1989 1999 2009 Summer condition 100 0 1959 180 14 Precipitation in late winter (Jan - March) p = 0.44, 6.6 cm increase over the 52 years Winter condition 160 200 Winter condition 2 Winter condition 3 Ice out date p = 0.003, 48 days later over the 52 220 Minimum Air Temperature in April p = 0.02, 2.1 degree increase over 52 years 12 Minimum Temperature 10 (All months are significantly different) 1959 - 1987 1995 - 2011 Temperature °C 8 6 4 2 0 -2 -4 -6 Jan 35 Feb Mar Apr May June July Aug Sept Oct Nov Dec Maximum Temperature * (* indicate significant change) * Temperature °C 30 25 * 20 15 10 5 0 Jan Feb Mar Apr May 45 July Aug Sept Oct Nov Dec Oct Nov Dec Precipitation 40 Centimeters of Precipitation June (* indicate significant change) 35 30 25 20 15 * 10 * 5 * 0 Jan Feb Mar Apr May June July Aug Sept Water temp August 0-5 meters p = 0.19, 0.33 degree increase over the 52 years 22 21 20 19 18 17 1959 5 4.5 1969 1979 1989 1999 Lake Temperature 23 2009 Shallow Primary productivity (0-12 m) p = 0.18, .74 increase over the 52 years 4 3.5 3 2.5 1 0.5 0 1959 1969 2.5 1979 1989 1999 2009 Deep Primary Productivity (13-32 m) p = 0.18, 0.4 decrease over the 52 years 2 1.5 1 0.5 0 1959 1969 1979 1989 1999 2009 Primary production 2 1.5 5 Carbon grams/square meter Shallow pooled mixed layerp = 0.0003 b pelagic PPr increases during the a drought and fish manipulation “experiment”, returning to a previous condition after the p = 0.045 experiment. Primary Productivity p = 0.0001 b p = 0.0045 4 3 a b a a a a 2 How much does climate a,bthe bamount of influence productivity during this experimental period? 1 0 pre exp pos t pre PPr 0 - 32 3500 pos t PPr 0 - 12 p = 0.0001 Fish Stocking b 2500 2000 p = 0.366 1500 a 1000 500 0 pre exp pos t Fi ngerl i ng pre exp Ca tcha bl e pre exp pos t PPr 13 - 32 b 3000 Pounds of fish stocked exp pos t pre exp PPr 0 - 5 pos t Outline • Lakes as sentinels of climate change – Physical and biological alterations linked to local and regional processes • Castle Station history: now the America’s oldest, continuous, mountain lake, ecological dataset • Long-term dynamics of climate, environmental, and biological variables • What is the influence of climate and fish manipulations on the zooplankton community composition? Primary Productivity 0 - 5 meters 5 Fish Experiment / drought period 4 3 2 1 1987 1988 1989 1990 1991 1992 1993 1994 1995 Carbon grams/square meter p = 0.0003 Primary Productivity 5 Shallow pooled mixed layer (0-5 m) pelagic b PPr increases during the drought andafish manipulation “experiment”, returning to a previous condition after the experiment. p = 0.045 p = 0.0001 b p = 0.0045 4 3 b a a a a a Total zooplankton biomass remains the a,bthere b is change in the functional same but players. 2 1 0 pre e xp pos t pre PPr 0 - 32 80 e xp pos t pre PPr 0 - 12 e xp pos t pre PPr 13 - 32 PPr 0 - 5 Zooplankton 70 p = 0.047 µg/liter 60 50 40 30 a p <0.0001 a b 20 10 b a,b b p <0.0001 p = 0.145 b p = 0.0002 b a a p = 0.1538 b b 0 pre exp pos t Di a ptomus pre exp pos t pre exp pos t pre exp pos t Cycl opoi d Da phni a Bos mi na pre exp pos t Hol opedi um e xp pre exp pos t Total pos t Zooplankton trends from an interannual viewpoint, not aggregated over period 35 Cyclopoid biomass p = 0.8, 7 increase over the 52 years 30 55 50 45 25 Daphnia biomass p = 0.06, 13 increase over the 52 years 40 35 20 30 25 15 20 10 15 10 5 5 0 1959 35 30 1969 1979 1989 1999 2009 0 1959 35 Diaptomus biomass p = 0.11, 12 decrease over the 52 years 30 25 20 20 15 15 10 10 5 5 0 1959 0 1959 1979 1989 1999 2009 1979 1989 1999 2009 1999 2009 Holopedium biomass p = 0.04, 8 decrease over the 52 years 25 1969 1969 1969 1979 1989 1959 - 1987 1995 - 2011 Summary of findings and future directions Long-term trends do not suggest changes in ice out, snow water content however, summer (min & max) and winter (min) air temperature is increasing. Shallow PPr is increasing but there are distinct periods of change; deep water PPr is decreasing. Climate does explain the change during the drought period as expected but the manipulation of fishes may cause an increase in PPr. Two decades are needed for the primary production to recover. Is this mediated through the zooplankton or nutrient excretion? Total zooplankton biomass has not changed over time. Functional players of zooplankton have shifted in the last 2 decades particularly in the copepod community which is dominated by cyclopoids. Fish are implicated in changing zooplankton community structure in recent times compared to the historical period likely due to the amount of stocking which seems to change the influence of lake variables on select taxa composition. Fish and climate (drought condition) together trump planktonic production and consumer composition, how long does it take to recover? Life history must matter? What about the influence of prior years on subsequent years? Ice out does not change but is something going on with nutrient cycling in the winter? Back to our conceptual model… now, we are focusing on maintaining our pelagic monitoring program but expanding to the collection of benthic production and invertebrate emergence, watershed inflow measurements for nutrients, winter sampling, fish analysis, linkages to terrestrial consumers (e.g. bats) Thanks for listening, come collaborate with us!