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Echoes in the ecosystem: top-down &
  bottom-up responses of Windermere to
       environmental perturbation

         Stephen Maberly, Ian Jones, Stephen
          Thackeray, Ian Winfield & Peter Henrys
                  Lake Ecosystem Group
      Centre for Ecology & Hydrology, Lancaster UK

NERC project in association with:
Jonathan Grey & Peter Smyntek- Queen Mary University of London
Project partners:
Mike Dobson- FBA
Chris Harrod- Queen’s University Belfast
Windermere
              • England’s largest lake
              • Two basins: deeper, less productive North
              and shallower more productive South
              • One of the most intensively studied lakes in
              the world
              • Long-term data and archives from early
              1900s and regular sampling for range of
              variables since 1945
              • Freshwater Biology Special Issue planned
              for Jan or Feb 2012




Photos from
FBA Image
Archive
Multiple stressors
        Natural variability in weather                     Schematic of changes in multiple
                                                              stressors in Windermere
                                                              Acidification                    Nutrient enrichment
                                                              Climate change                   Invasion of non-native species

                                                                                   Tertiary           Detectable
                             Climate change            Sew age w orks
                                                       installed
                                                                        Sulfate
                                                                        loading    treatment          change in w ater
    Acid                                                                declines                      temperature


(& nutrients)                                                                                                   Large No.s of
                                                       1                                                        roach detected

                Nutrients
                (& toxins)                         0.8

                                                   0.6
                                              Stress
                                                   0.4

                                                   0.2

                                                       0
                                                       1940 1950 1960 1970 1980 1990 2000 2010
                                                                         Year
Publications on invasive species & freshwater




                 Search on Web of Science, May 2011
Response to changing P
40
     Mean winter TP (mg m-3)                                                                                               Mean Chla (mg m )
                                                                                                                                            -3

35                                                                                                         12

30                                                                                                         10

25
                                                                                                            8
20
                                                                                                            6
15

10                                                                                                          4
        North Basin
 5                                                                                                          2         North Basin
        South Basin
 0                                                                                                                    South Basin
                                                                                                            0
 1950   1960   1970      1980   1990        2000   2010
                                                                                                            1950    1960     1970   1980   1990   2000   2010
                         Year
                                                                                                                                    Year
                                                                                                           10
                                       -3                                                                               Min O2 at depth (g m-3)
        Mean winter SRP (mg m )
30                                                                              Max pH

                                                          10
                                                                                                            8
25         North Basin
           South Basin
20                                                                                                          6
                                                           9
15
                                                                                                            4

10                                                         8
                                                                                                            2
                                                                  North Basin                                       North Basin
5
                                                                  South Basin                                       South Basin
                                                           7                                                 0
0
                                                           1950   1960   1970     1980    1990   2000   2010 1950   1960     1970   1980   1990   2000   2010
1950    1960   1970      1980   1990        2000   2010                            Year                                             Year
                         Year
Seasonal patterns of phytoplankton change

                                 Spring                                                                         Summer

                         20          North Basin      South Basin                                          20


                         16                                                                                16
Chla in M,A,M (mg m-3)




                                                                                  Chla in J,J,A (mg m-3)
                         12                                                                                12


                          8                                                                                 8


                          4                                                                                 4


                          0                                                                                 0
                          1960      1970     1980          1990     2000   2010                             1960   1970   1980          1990   2000   2010
                                                    Year                                                                         Year

                                                                      Lines are 3-year moving averages
Zooplankton
                                                                        40       North Basin
18                             -3
       Mean zooplankton (No. dm )                                                                1977-1991
16




                                           Mean Zooplankton (No. dm )
                                                                                                 1993-2007




                                                                   -3
14
                                                                        30
12

10

8                                                                       20
6

4         North Basin
2         South Basin                                                   10
0
1950           1970     1990        2010

                                                                         0
                                                                             0      3     6        9     12
                                                                                         Month
Fish predation?
                          6000




                                                                         CPUE (fish (Fish 100 m net-1day ) -1)
                                                                                                                 50




                                                                              CPUE 100 m net-1 day
                                                                                         -1
Abundance (fish ha )-1)




                          5000
                                                                                                                 40
   Abundance (fish ha
         -1




                          4000
                                                                                                                 30




                                                                                         2
                                                                                              2
                          3000                                                                                   20


                          2000                                                                                   10


                          1000                                                                                   0
                                                                                                                      1995   2000          2005   2010
                                                                                                                                    Year
                            0
                             1990   1995      2000         2005   2010
                                               Year

                                           Largely roach
Climate change and roach invasion?
                                                                           Maitland 1972
                                                 Roach
                                                 distribution
     Mean surface temperature (oC)

12



10
                                                                         Davies et al. 2004

8
           North Basin
           South Basin

6                                       ‘It is not unlikely that these
                                        had been brought as live-
 1950       1970          1990   2010   bait for pike, as live-baiting
                   Year                 is occasionally done by
                                        strangers.’
                                        (Watson, 1925)
Possible consequences for Arctic charr
Catch per unit effort on spawning grounds




                                            100                                                                                              100                                              20
                                                             North Basin                                                                                     North Basin




                                                                                                 Catch per unit effort on spawning grounds
                                            90                                                                                               90                                               18




                                                                                                                                                                                                   Summer zooplankton (No. dm-3)
                                            80                                                                                               80                                               16
                                            70                                                                                               70                                               14
                                            60                                                                                               60                                               12
                                            50                                                                                               50                                               10
                                            40                                                                                               40                                               8
                                            30                                                                                               30                                               6
                                            20                                                                                               20                                               4
                                                                                                                                                        Arctic charr
                                            10             Arctic charr                                                                      10                                               2
                                                                                                                                                        Summer zooplankton
                                             0                                                                                                0                                               0
                                             1950   1960   1970      1980   1990   2000   2010                                                1950   1960   1970   1980      1990   2000   2010
                                                                     Year                                                                                          Year
• Echoes in the ecosystem
Carnivores

                                         Changes in
                                          Pike diet
Planktivores     Increase in                                         Reduction in
                    roach                                            Arctic charr

Zooplankton                      Reduction in
                                 zooplankton


                                        Increase in
Phytoplankton                         phytoplankton



 Chemistry                                              Increased     Reduction
                                                       internal P-   in oxygen at
    Climate                                                load          depth
    change
                        Warmer               Stronger
                         water             stratification               Physics
Mixed layer depth & light availability
                             Day of year
             0
                  50   70   90      110    130   150
             -5

            -10

            -15
                                                       seventies
Depth (m)




            -20                                        eighties

            -25                                        nineties
                                                       naughties
            -30

            -35

            -40




                                                                   seventies


                                                                               naughties
            -45



                            South Basin
Phytoplankton edibility changes?
                                                                                                                                1000000000
               1000000000




                                                                                   Biolvol in JJA of algae < or >50 µm (µm3 cm-3)
                                                     >50 um
Average biovolume in JJA (µm3 cm-3)




                                                                                                                                    100000000
                             100000000                                                                                                                     y = 1E+07e-0.152x   P<0.05
                                                     <50 um                                                                                                   R² = 0.155



                                      10000000
                                                                                                                                    10000000



                                       1000000

                                                                                                                                     1000000

                                       100000


                                                                                                                                      100000
                                         10000

                                                                                                                                                    y = 306565e-0.065x
                                                                                                                                                        R² = 0.174     P<0.05
                                         1000                                                                                          10000
                                             1950      1970          1990   2010                                                                0      5        10         15        20

                                                              Year                                                                          Zooplankton density in JJA (No.       dm-3)

                                                                                                                                                                      North Basin
Path-analysis for the North Basin (Bayesian belief
         network implemented in Winbugs)


      Roach                                      Arctic charr
     numbers                                      numbers

              4%
                   Zooplankton
                    density in
                     summer
                          12%

                                 Phytoplankton
                                    (Chla) in
                                    summer

                                                    Oxygen
                                                 concentration
   Water                                           at depth
temperature                                              6%
        30%
Conclusions

• Multiple stressors have affected Windermere over the
  last 65 years via top-down and bottom-up processes- the
  response of a complicated ecosystem to perturbation is
  complex
• Nutrient enrichment has had the major impact but more
  recently climate change interacting with expansion of a
  non-native species has started to alter the food-web and
  the way the lakes converts resources into algae
• Climate change is likely to make the achievement of
  water quality and conservation objectives more difficult
  and nutrient targets will have to be more stringent
Acknowledgements
• We thank the FBA (1945 to 1989) and our
  colleagues in CEH/IFE (1989-present) for
                         (1989-
  collecting the long-term data analysed here
                 long-

• This work was funded by a NERC EHFI grant to
  CEH and QMUL



• See our web-page: http://www.windermere-
          web-      http://www.windermere-
  science.org.uk/home

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Echoes in the Ecosystem

  • 1. Echoes in the ecosystem: top-down & bottom-up responses of Windermere to environmental perturbation Stephen Maberly, Ian Jones, Stephen Thackeray, Ian Winfield & Peter Henrys Lake Ecosystem Group Centre for Ecology & Hydrology, Lancaster UK NERC project in association with: Jonathan Grey & Peter Smyntek- Queen Mary University of London Project partners: Mike Dobson- FBA Chris Harrod- Queen’s University Belfast
  • 2. Windermere • England’s largest lake • Two basins: deeper, less productive North and shallower more productive South • One of the most intensively studied lakes in the world • Long-term data and archives from early 1900s and regular sampling for range of variables since 1945 • Freshwater Biology Special Issue planned for Jan or Feb 2012 Photos from FBA Image Archive
  • 3. Multiple stressors Natural variability in weather Schematic of changes in multiple stressors in Windermere Acidification Nutrient enrichment Climate change Invasion of non-native species Tertiary Detectable Climate change Sew age w orks installed Sulfate loading treatment change in w ater Acid declines temperature (& nutrients) Large No.s of 1 roach detected Nutrients (& toxins) 0.8 0.6 Stress 0.4 0.2 0 1940 1950 1960 1970 1980 1990 2000 2010 Year
  • 4. Publications on invasive species & freshwater Search on Web of Science, May 2011
  • 5. Response to changing P 40 Mean winter TP (mg m-3) Mean Chla (mg m ) -3 35 12 30 10 25 8 20 6 15 10 4 North Basin 5 2 North Basin South Basin 0 South Basin 0 1950 1960 1970 1980 1990 2000 2010 1950 1960 1970 1980 1990 2000 2010 Year Year 10 -3 Min O2 at depth (g m-3) Mean winter SRP (mg m ) 30 Max pH 10 8 25 North Basin South Basin 20 6 9 15 4 10 8 2 North Basin North Basin 5 South Basin South Basin 7 0 0 1950 1960 1970 1980 1990 2000 2010 1950 1960 1970 1980 1990 2000 2010 1950 1960 1970 1980 1990 2000 2010 Year Year Year
  • 6. Seasonal patterns of phytoplankton change Spring Summer 20 North Basin South Basin 20 16 16 Chla in M,A,M (mg m-3) Chla in J,J,A (mg m-3) 12 12 8 8 4 4 0 0 1960 1970 1980 1990 2000 2010 1960 1970 1980 1990 2000 2010 Year Year Lines are 3-year moving averages
  • 7. Zooplankton 40 North Basin 18 -3 Mean zooplankton (No. dm ) 1977-1991 16 Mean Zooplankton (No. dm ) 1993-2007 -3 14 30 12 10 8 20 6 4 North Basin 2 South Basin 10 0 1950 1970 1990 2010 0 0 3 6 9 12 Month
  • 8. Fish predation? 6000 CPUE (fish (Fish 100 m net-1day ) -1) 50 CPUE 100 m net-1 day -1 Abundance (fish ha )-1) 5000 40 Abundance (fish ha -1 4000 30 2 2 3000 20 2000 10 1000 0 1995 2000 2005 2010 Year 0 1990 1995 2000 2005 2010 Year Largely roach
  • 9. Climate change and roach invasion? Maitland 1972 Roach distribution Mean surface temperature (oC) 12 10 Davies et al. 2004 8 North Basin South Basin 6 ‘It is not unlikely that these had been brought as live- 1950 1970 1990 2010 bait for pike, as live-baiting Year is occasionally done by strangers.’ (Watson, 1925)
  • 10. Possible consequences for Arctic charr Catch per unit effort on spawning grounds 100 100 20 North Basin North Basin Catch per unit effort on spawning grounds 90 90 18 Summer zooplankton (No. dm-3) 80 80 16 70 70 14 60 60 12 50 50 10 40 40 8 30 30 6 20 20 4 Arctic charr 10 Arctic charr 10 2 Summer zooplankton 0 0 0 1950 1960 1970 1980 1990 2000 2010 1950 1960 1970 1980 1990 2000 2010 Year Year
  • 11. • Echoes in the ecosystem Carnivores Changes in Pike diet Planktivores Increase in Reduction in roach Arctic charr Zooplankton Reduction in zooplankton Increase in Phytoplankton phytoplankton Chemistry Increased Reduction internal P- in oxygen at Climate load depth change Warmer Stronger water stratification Physics
  • 12. Mixed layer depth & light availability Day of year 0 50 70 90 110 130 150 -5 -10 -15 seventies Depth (m) -20 eighties -25 nineties naughties -30 -35 -40 seventies naughties -45 South Basin
  • 13. Phytoplankton edibility changes? 1000000000 1000000000 Biolvol in JJA of algae < or >50 µm (µm3 cm-3) >50 um Average biovolume in JJA (µm3 cm-3) 100000000 100000000 y = 1E+07e-0.152x P<0.05 <50 um R² = 0.155 10000000 10000000 1000000 1000000 100000 100000 10000 y = 306565e-0.065x R² = 0.174 P<0.05 1000 10000 1950 1970 1990 2010 0 5 10 15 20 Year Zooplankton density in JJA (No. dm-3) North Basin
  • 14. Path-analysis for the North Basin (Bayesian belief network implemented in Winbugs) Roach Arctic charr numbers numbers 4% Zooplankton density in summer 12% Phytoplankton (Chla) in summer Oxygen concentration Water at depth temperature 6% 30%
  • 15. Conclusions • Multiple stressors have affected Windermere over the last 65 years via top-down and bottom-up processes- the response of a complicated ecosystem to perturbation is complex • Nutrient enrichment has had the major impact but more recently climate change interacting with expansion of a non-native species has started to alter the food-web and the way the lakes converts resources into algae • Climate change is likely to make the achievement of water quality and conservation objectives more difficult and nutrient targets will have to be more stringent
  • 16. Acknowledgements • We thank the FBA (1945 to 1989) and our colleagues in CEH/IFE (1989-present) for (1989- collecting the long-term data analysed here long- • This work was funded by a NERC EHFI grant to CEH and QMUL • See our web-page: http://www.windermere- web- http://www.windermere- science.org.uk/home