The Influence of Natural and Anthropogenic Perturbations on Lake Riparian Forest and Coarse Woody Debris Modeling Linkages Between Aquatic and Terrestrial Ecosystems September 26, 2002 Greg Sass This is a collaborative effort! • • • • NSF-Biocomplexity Project Dr. Monica Turner Dr. Stephen Carpenter Isaac Kaplan, Anna Sugden-Newbery, Anthony Yannarell, Theodore Willis, Greg Sass • Scott van Egeren, Michelle Parara Biocomplexity Riparian forest, land, people, and lakes http://limnology.wisc.edu Click on research link, follow to biocomplexity web page Relationship between CWD density and shoreline development in N. Wisconsin lakes -Also true for MN macrophytes! CWD Density (no./km) Shoreline Development From Christensen et al. 1996 Relationship between fish growth and coarse woody debris (CWD) in N. Wisconsin lakes Undeveloped Undeveloped log Growth Rate (mm/yr) Low Development Low Development High Development High Development CWD Density (no./km) From Schindler et al. 2000 How are changes on land and land/water interface reflected in the adjacent lake ecosystem? • Does the riparian forest influence fish populations? – The riparian forest is linked to fish through Coarse Woody Debris (CWD) CWD abundance influenced by: • Forest structure (Harmon et al. 1986, Hely et al. 2000) – Successional state • Natural and anthropogenic disturbance (Christensen et al. 1996, Guyette and Cole 1999 , Hely et al 2000) – Windthrows – Logging – Lakeshore development Photo courtesy of Michelle Parara Why model CWD dynamics? • These are big systems with slow (and fast) dynamics!! Photo courtesy of Michelle Parara 923 yearold white pine in Swan Lake, Ontario Guyette and Cole 1999 Three Aspects Compose the Linked Terrestrial-Aquatic Model Terrestrial Terrestrial-Aquatic Interface Riparian Model Aquatic Fish Model Main Goals of the Wood Model 1. Create CWD via riparian forest that can be affected by both natural and anthropogenic processes 2. Simulate realistic CWD densities that can be used to test hypotheses/ask questions about effect of CWD on fish communities Conceptual Structure of the Wood Model Recruitment Saplings Falling away from water Graduation Adults Trees that die and stay upright Trees that die and fall immediately Snags Falling away from water CWD Falling Loss to decay and deep water Two Pools of Trees: • “Softwoods” – Representative of early succession canopy – Paper birch (Betula papyrifera), aspens (Populus spp) Big tooth aspen (Populus grandidentata) Paper birch (Betula papyrifera) • “Hardwoods” – Representative of mid-late succession canopy – White pine (Pinus strobus), sugar maple (Acer saccharum) White pine (Pinus strobus) Sugar maple (Acer saccharum) Riparian Model Formulas • SAPLINGS “Shading” terms Si(t+1) = Si(t) + {Ai(t)ri * (1-αjiAj(t) – αjiAi(t))} - Si(t)gi • ADULT TREES Ai(t+1) = Ai(t) + Si(t)gi - mi Ai(t); • STANDING SNAGS SSi(t+1) = SSi(t) + (1-Li) mi Ai(t) - fi SSi(t); • COARSE WOODY DEBRIS Di(t+1) = Di(t) + (γ fi SSi(t)) + {γ fi SSi(t) Li mi Ai(t)} - (a11 + a21) Di(t) ; Conceptual Structure of the Wood Model (+) Hardwood Saplings “Shading” (-) Graduation Recruitment Adult Hardwoods Softwood Saplings (+) (-) Graduation Adult Softwoods Recruitment Model Scenarios • Baseline Scenario – Riparian forest density from Turner 2001 and Christensen et al. 1996 – CWD values from undeveloped Little Rock Lake in Vilas County, WI • Windthrow Scenario – 65% instantaneous death of hardwood and softwood adults and snags • Clearcut Scenario – 95% instantaneous death of hardwood and softwood adults and snags Little Rock Lake • Development Scenario – 1% annual loss of adult hardwoods and softwoods – 5% annual loss of Snags and CWD Adult Tree and CWD Dynamics During Baseline Scenario # Logs/km shoreline # Trees/km Shoreline Hardwoods Softwoods 2000 1500 Adult Trees 1000 500 0 600 400 CWD 200 0 0 50 100 150 200 250 300 Years 350 400 450 500 Adult Tree and CWD Dynamics During Windthrow Scenario # Logs/km shoreline # Trees/km Shoreline Hardwoods Softwoods 2000 1500 Adult Trees 1000 500 0 600 400 CWD 200 0 0 50 100 150 200 250 Years 300 350 400 450 500 CWD Dynamics During Windthrow Scenario Hardwood + Softwood CWD/km Shoreline 800 600 400 200 0 0 100 200 300 Years 400 500 CWD Abundance (all Trees) Following FireDisturbance Hely et al. 2000 Adult Tree and CWD Dynamics During Clearcut Scenario # Logs/km Shoreline # Trees/km Shoreline Hardwoods Softwoods 2000 1500 Adult Trees 1000 500 0 600 400 CWD 200 0 0 100 200 300 Time (years) 400 500 Hardwood + Softwood CWD/km Shoreline CWD Dynamics During Clearcut Scenario 2500 2000 1500 1000 500 0 0 100 200 300 Time (years) 400 500 Adult Tree and CWD Dynamics During Development Scenario # Trees/km Shoreline 2000 # Logs/km shoreline Hardwoods Softwoods 600 1500 Adult Trees 1000 500 0 400 CWD 200 0 0 50 100 150 200 250 300 Years 350 400 450 500 CWD Dynamics during Development Scenario Hardwood + Softwood CWD/km Shoreline 800 600 400 200 0 0 50 100 150 200 250 300 350 400 450 500 Years Can the model mimic ‘real’ history? Skidding Logs, Upper Chippewa Basin, Circa 1890 Lakeshore Development Last ~50 years # Logs/km Shoreline # Trees/km Shoreline Taming the Northwoods Hardwoods Softwoods 2000 Clear cut Development 1500 Adult Trees 1000 500 0 600 400 CWD 200 0 0 100 200 300 Time (years) 400 500 CWD Dynamics in Clearcut + Development Scenario # Logs/km Shoreline 600 400 200 0 0 100 200 300 Time (years) 400 500 Summary of Wood Model • Model is simple, but fairly realistic • Windthrows and clearcuts have long-term effects on CWD pool • Development a powerful force Conclusions • This ecosystem-level model is a useful tool for creating questions about CWD inputs/removals. – Can we devise ways to observe long-term changes in riparian forest and CWD structure? – Does indiscriminate thinning actually occur? – How long does it take for the CWD pool to recover? – How do changes in CWD abundance affect fish communities? • How can we obtain answers to these questions? Biocomplexity Cross-lakes Crew • Led by Michelle Parara and Scott van Egeren • Riparian forest/CWD analysis: Anna Sugden-Newbery Modeling linkages between terrestrial and aquatic ecosystems part II: The influence of riparian forest dynamics on aquatic food webs Isaac Kaplan, Tanya Havlicek, Pieter Johnson, Brian Roth,Greg Sass, Anna Sugden-Newbery, Theodore Willis, Anthony Yannarell, Monica Turner, and Steve Carpenter Biocomplexity Riparian forest, land, people, and lakes Relationship between fish growth and coarse woody debris in N. Wisconsin lakes Undeveloped Undeveloped log Growth Rate (mm/yr) Low Development Low Development High Development High Development Coarse Woody Debris Density (no./km) From Schindler et al. 2000 Conceptual Model Terrestrial Terrestrial-Aquatic interface Aquatic Growth Senescence coarse woody debris Forest Aquatic Food Web Windthrow Humans Development Decay/ Physical Transport Fishing Questions • How does the aquatic food web respond to stable levels of coarse woody debris? • How does the food web respond to perturbations? - windthrow, development, fishing • How can we learn about effects of coarse woody debris on fish predation and growth rates in experimental lakes? Fish biomass dynamics model adult piscivore juv. piscivore benthivore insects Hypothesized Effects of Coarse Woody Debris on Fish Community Insect Abundance kg of trichopterans + odonates /ha 1 0.8 0.6 0.4 0.2 0 90 0 75 0 60 0 45 0 30 0 15 0 0 logs /km shoreline Response of Vulnerability and Hiding parameter value 2 vulnerability of benthivore 1.5 hiding of benthivore 1 vulnerability of juv. piscivore 0.5 hiding of juv. piscivore 0 0 200 400 600 800 logs/km of shoreline Fish Model: Benthivore Biomass Equation dB/dt=G-mB2-P2 –P3 G=fishC*g1 +BugC*g2 Bt+1=Bt-harvest --------------------------------------------------Functional Response: Piscivory v1 B1-V1 hiding V1 h1 vulnerable c12 B2 predators Hypothesis 1: Similar piscivore and benthivore behavioral response to logs Response of Vulnerability and Hiding parameter value 2 vulnerability of benthivore 1.5 hiding of benthivore 1 vulnerability of juv. piscivore 0.5 hiding of juv. piscivore 0 0 200 400 600 800 logs/km of shoreline kg/ hectare Fish Biomass at Steady State 35 30 25 20 15 10 5 0 benthivore juv. piscivore adult piscivore 50 500 1000 logs / km of shoreline 500 and fishing Hypothesis 2: Benthivore is less dependent on refuge than piscivore Response of Vulnerability and Hiding parameter value 2 vulnerability of benthivore 1.5 hiding of benthivore 1 vulnerability of juv. piscivore 0.5 hiding of juv. piscivore 0 0 100 200 300 400 logs/km of shoreline Fish Biomass at Steady State kg/ hectare 50 40 benthivore 30 juv. piscivore 20 adult piscivore 10 0 50 500 1000 logs / km of shoreline 500 and fishing Fish Biomass Response to Windthrow Windthrow 30 600 500 20 400 300 10 200 100 0 100 Fish Biomass Response to Windthrow and Fishing 200 300 400 0 500 years 30 600 500 20 400 300 10 200 100 0 100 200 300 years 400 0 500 fishing starts Logs /km of shoreline kg of fish / hectare 700 Benthivore Juvenile Piscivore Adult Piscivore Coarse Woody Debris Logs /km of shoreline kg of fish / hectare 700 Fish Biomass Response to Development Development 30 500 400 20 300 200 10 100 0 100 Fish Biomass Response to Development and Fishing 200 300 400 0 500 years 30 500 400 20 300 200 10 100 0 100 200 300 years 400 0 500 fishing starts Logs /km of shoreline kg of fish / hectare 600 Benthivore Juvenile Piscivore Adult Piscivore Coarse Woody Debris Logs /km of shoreline kg of fish / hectare 600 Conclusions • Coarse woody debris could be a major driver of fish community dynamics (and we will test this) • Effect of development is much greater than effect of windthrow • For benthivore, moderate reductions in coarse woody debris may be balanced by fishing on piscivore • Help Greg chop down trees this winter Work in Progress Field experiments in N. Wisconsin: • Removal of coarse woody debris from Little Rock Lake • Addition of coarse woody debris to Camp Lake • Observations of growth and abundance • Estimation of predation and vulnerability parameters and hypothesis testing Acknowledgements Michele Parara, Scott VanEgeren, and the Biocomplexity Field Crew This work is funded by the National Science Foundation under Cooperative Agreement #DEB-0083545 Piscivore Response to CWD Benthivore Response to CWD 60 kg of fish /ha kg of fish /ha 40 30 40 20 20 10 0 1000 0 1000 500 logs / km 0 0 50 100 150 200 300 250 inflection pt. for benthivore 500 200 logs / km 0 0 100 inflection pt. for benthivore Increasing vulnerability Increasing vulnerability of benthivore of benthivore