1. Water Dynamics: The Experimental Perspective Alan K. Knapp Graduate Degree Program in Ecology, Colorado State University

2. Overview: Experimental approach to: Climate Change - Ecosystem Function (in grasslands) Soil moisture & ANPP (4 studies) Change in precipitation pattern (long-term) Comparison among grasslands (short-term)

3. Sala et al. 1988 Rainfall vs. Production across the Central US Grasslands and similar ecosystem types cover ~ 40% of the Earth’s land surface Most are water limited systems Most of what we know is based on observation Rainfall regimes CO 2 effects Temperature effects Soil moisture + + - - Organism Water relations Ecosystem Process & Function

4. Soil moisture Inputs Outputs N C

5. Central US grasslands provide a template for experiments at sites that span key precipitation and temperature gradients… “Seeing the grasslands beyond the grasses”

6. Mesic Grasslands of North America The Flint Hills of Kansas – The Konza Prairie Tallgrass prairie is the most mesic of US Great Plains grasslands – sufficient precipitation to support forest Flint Hills are located at the dry edge of the original range and on shallow soils KPBS C 4 grass dominance A dynamic & tractable system

7. Proportion of total rainfall in the US from large (>5 cm) rainfall events Proportion (%) of rainfall in “extreme events” Abundant evidence that rainfall regimes are changing Oklahoma Question: (We know amount is important) Given increased variability and frequency of extreme events, to what extent will altered precipitation patterns affect ecological processes in “mesic” grasslands?

8.  12 Rainfall Manipulation Plots (RaMPs) + non-sheltered controls  Collect, store, and reapply natural rainfall on intact prairie plots  Treatments include ambient and altered rainfall patterns since 1998  Elevated temperature treatment added in 2003 The Rainfall Manipulation Plot (RaMP) Experiment Address the impact of changes in size and timing of growing season rainfall events and elevated temperature

9. Target altered rainfall pattern: 50% longer inter-rainfall periods, larger individual rain events No change in total precipitation quantity Fewer events Greater storm size Increased dry periods

10. Ambient ppt timing: “typical” seasonal pattern Altered ppt timing: repeated deficits more extreme wetting and drying cycles Soil moisture dynamics Average soil water content in top 30 cm: - reduced by 12% Variability in soil moisture: - increased by 27%

11. ANPP responses to altered rainfall timing 12-22% reduction in years with significant responses 11% reduction across years (p=0.01)

12. Across all years and all rainfall treatments

13. If only the entire world worked like Konza…. Can we apply results from mesic grasslands to other climates and grasslands?

14. Ojima and Lackett 2002 Tallgrass Mixed-grass Short-grass Precipitation gradient A multi-system assessment of climate change responses Heisler-White et al. 2009. Global Change Biology.

15. Semi-arid grassland Extreme rainfall events, long dry intervals = reduced ANPP? With the same experimental design, how will these different grasslands respond? Mesic grassland Extreme rainfall events, long dry intervals = reduces ANPP

16. SGS Experimental manipulation of growing season rainfall regimes Frequent, small events vs. infrequent large events Equal rainfall amounts within each site A = 12 events B = 6 events C = 4 events Total = 191 mm A = 12 events B = 6 events C = 4 events Total = 450 mm KNZ Semi-arid Mesic

17. 15% decrease in ANPP with fewer large events 30% increase in ANPP with fewer large events 191 mm 340 mm 450 mm 70% increase in ANPP with fewer large events Semi-arid steppe Mixed grass prairie Tallgrass prairie Heisler-White et al. 2009

18. Why do arid vs. mesic grasslands respond differently? Two key aspects of more extreme rainfall regimes: Fewer events Greater storm size Increased dry periods 1. Larger rain events 2. Longer dry periods between events When soils are usually moist, longer dry periods negatively impact ANPP… When soil are usually dry, larger rain events improve soil water and increase in ANPP…

19. The Sevilleta LTER – the edge of the Great Plains

20. Monsoon Rainfall Manipulation Experiment During the summer monsoon July through September: 3 plots receive ambient rainfall 5 plots receive a 5mm rainfall event every week 5 plots receive a 20 mm rainfall event every month Scott Collins Michell Thomey Rodrigo Vargas Jennifer Johnson Renee Brown Don Natvig Mike Friggens

21. Seasonal average soil water content, predawn water potentials, leaf-level A net and soil CO 2 flux were highest in plots receiving a few large events Thomey et al. 2011 GCB Responses similar to shortgrass steppe, but opposite mesic grasslands

22. Total ANPP and ANPP of black grama were highest in plots receiving a few large events Thomey et al. 2011 GCB 25 – 40% greater ANPP of black grama with few large events compared to many small events Similar trends for ANPP of all species combined

23. Responses to drought - is there a pattern/amount interaction? Mixed- grass Shortgrass Tallgrass SGS-LTER (North) Fort Union (South) Sand Creek (Central) Karie Cherwin

24. From Huxman et al (2004) These are observational data!

25. Testing this inference of high sensitivity 30 plots at each site –10 controls –20 rainout shelters 10 at 50% Reduced- PPT 10 at 80% Reduced- PPT Two-year drought Response Variables –Net Primary Productivity –Soil Moisture No change in timing/pattern Reduces event size & total amount

26. a b c a b c a b c a b c a b c a b c South Central North

27. CentralSouth a a b b b b x x y

28. Sensitivity to Drought 40° N35° N38° NLatitude: Temperature: Precipitation: Elevation: Soil Texture: 7.2° C Coarse 8.3° C9.4° C 310 - 330 mm410 - 450 mm330 - 360 mm 1650 m2043 m1219 m FineIntermediate (Sandy Loam) (Clay) (Sandy Clay Loam) 2007 2008 Low High Why the differential sensitivity? Insensitive ?!?! Within biome saturation?

29. Rainfall regimes – there’s more to them than amount… In addition to more rain, wet years have more large events compared to average years.

30. 33.6 events 41.0 events 9.7 mm 5.3 mm Insensitive Sensitive Many, small events (dry, low ANPP) Fewer, large events (wet, high ANPP)

31. What does all this mean?!?! There is still much we need to learn about precipitation regimes, soil moisture and ecosystem responses if we are to forecast the future… Because all three are likely to change. Much of what we know is from observation and correlation… We need to challenge this knowledge Alterations in rainfall patterns (and of course amounts) may change productivity in biome specific ways… Alterations in rainfall patterns may even alter sensitivity to drought…

32. As Ecologists – one of our greatest challenges is to provide knowledge to help us forecast and cope with future ecosystem structure and function… The empirical perspective - future field experiments? We need more long-term experiments where we “push” ecosystems… in both magnitude and pattern… We need comparative analyses of ecosystem responses to climate change within and across biomes… Clear need for long-term, multi-site and coordinated experiments… and these should be designed with modelers…

33. Divergent responses are linked to differential effects of extreme rainfall events on mean soil water content ↑ SWC by 19% in arid site ↓ SWC by 20% in mesic site

34. ResponsePredictor variablePartial r 2 Model r 2 Total ANPPCV SWC 15cm0.598 (92%)0.651 mean SWC 15cm0.053 Grass ANPPmean SWC 15cm0.470 (70%)0.676 mean SWC 30cm0.103 CV SWC 30cm0.103 Forb ANPPmean SWC 30cm0.375 (82%)0.453 CV SWC 30cm0.078 Grasses and forbs respond to different aspects of soil water dynamics Grasses most responsive to mean surface SWC, while forbs respond more to deeper SWC Potential for long-term changes in community composition