Differential impacts of Pleistocene mega-herbivores on nitrogen availability

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1 Differential impacts of Pleistocene mega-herbivores on nitrogen availability Elizabeth S. Jeffers Long Term Ecology Laboratory Department of Zoology University of Oxford

2 Extinctions were coincident with major other ecological changes Source: Willerslev, E., et al. (2014) Nature 506(7486): Source: Lorenzen, E. et al. (2011). Nature 479(7373): Source: McLauchlan, K. K., et al. (2013). Nature 495:

3 What were the causes and consequences of past extinctions? Climate Solar Radiation Atmospheric Geochemistry Human (over)predation Change in plant species composition & diversity Plant biomass Woody species, Herbs Nutrient cycling Change in herbivore assemblages Extinction of large herbivores Plant biomass Woody species Herbs Nutrient cycling Change in plant species composition & diversity

4 How to tease apart causes & consequences? Regional to Global Abiotic Factors Local to Regional Biotic Factors Local Abiotic Factors Mechanistic Modelling of palaeo-population dynamics = 1 = 1 1 P

5 Late-Glacial British and Irish Ecosystem Dynamics Regional to Global Abiotic Factors Local to Regional Biotic Factors Local Abiotic Factors

6 Late-Glacial British and Irish Ecosystem Dynamics Lough Nadourcan, Ireland Quidenham Mere, England Empetrum heathland to birch woodland Large Herbivore Biomass following above ground biomass shrubs & arrival of first trees N availability with above ground plant biomass Grassland to deciduous woodland Large Herbivore Biomass following above ground biomass shrubs & arrival of first trees N availability with above ground plant biomass Jeffers, E. S., M. B. Bonsall, et al. (2012). New Phytologist 193(1): Jeffers, E. S., N.J. Whitehouse, et al. (in prep.). Jeffers, E. S., M. B. Bonsall, et al. (2011). Journal of Ecology 99(4):

7 Mechanistic Modelling of Ecosystem Change - Lough Nadourcan More Betula(birch) biomass consumed by large herbivores than Empetrum(crowberry) More Betula(birch) biomass burned than Empetrum (crowberry) Betula(birch) takes up more N than Empetrum (crowberry) for equivalent rise in aboveground biomass Jeffers, E. S., M. B. Bonsall, et al. (2012). New Phytologist193(1):

8 Mechanistic Modelling of Ecosystem Change Lough Nadourcan Rank What Drivers Most Important? Betula Rank Empetrum 1 Fire Activity 1 Herbivore Biomass 2 Growing Season Temperature 2 Fire Activity 3 Herbivore Biomass 3 Growing Season Temperature 4 N availability 4 N availability Jeffers, E. S., M. B. Bonsall, et al. (2012). "Climate change impacts on ecosystem functioning: evidence from an Empetrum heathland." New Phytologist 193(1):

9 Mechanistic Modelling of Ecosystem Change Quidenham Mere Which breakpoint best describes changes through time in oak-grass competition? Oak Dominance Low Herbivore Biomass Low Fire Activity Warmer summer temperatures Grass & Herb Dominance High Herbivore Biomass High Fire Activity Cool summer temperatures AIC weight 5% 95% Declining N availability Increasing N availability 0% 7500 cal yrsbp Time cal yrs. BP Jeffers, E. S., M. B. Bonsall, et al. (2011). Journal of Ecology 99(4):

10 Impact of Large Herbivores on N availability Lough Nadourcan Jeffers, E. S., M. B. Bonsall, et al. (2012). ; Jeffers, E.S., N.J. Whitehouse, et al (in prep)

11 Impact of Large Herbivores on N availability Quidenham Mere Jeffers, E. S., M. B. Bonsall, et al. (2011). Jeffers E.S., N.J. Whitehouse, et al. (in prep)

12 Conclusions Guild-specific response of large herbivores to woody plant encroachment Climate and fire direct drivers of tree expansion, herbivores had indirect effects Impact of herbivores on terrestrial N availability depends on herbivore assemblage, plant species composition, geology

13 Acknowledgements Adrian Lister (NHM, London) Nicki Whitehouse (Plymouth) Steve Brooks (NHM,London) Kendra McLauchlan (Kansas State) Kathy Willis (Oxford) Mike Bonsall(Oxford) = 1 Members of the Oxford Long-Term Ecology Laboratory

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