Western Mountain Initiative The climatic water balance in an ecological context Nathan L. Stephenson U.S. Department of the Interior U.S. Geological Survey Thesis: Consideration of the climatic water balance improves our ability to understand and forecast the effects of climatic changes on the biological world. (1) The superiority of the climatic water balance in ecological studies. (2) In a changing climate, increasing water availability will NOT counteract the effects of increasing evaporative demand. … And why soil moisture may be less important than you think! (3) Don’t forget AET! Thesis: Consideration of the climatic water balance improves our ability to understand and forecast the effects of climatic changes on the biological world. (1) The superiority of the climatic water balance in ecological studies. (2) In a changing climate, increasing water availability will NOT counteract the effects of increasing evaporative demand. … And why soil moisture may be less important than you think! (3) Don’t forget AET! Mean annual temperature (C) Mean annual precipitation (cm) Whittaker 1975 Traditional view: At broad scales, vegetation type is a function of some measure of annual energy supply, annual water supply, or their ratios or differences. Mean annual temperature (C) Temperature Potential evapotranspiration (PET) Net radiation Growing degree-days Precipitation (P) P/PET (or PET/P) PET-P (or P-PET) Mean annual precipitation (cm) Whittaker 1975 Traditional view: At broad scales, vegetation type is a function of some measure of annual energy supply, annual water supply, or their ratios or differences. The Big Problem: The traditional view does not consider the seasonal interactions of energy and water. E.g., with nearly identical mean annual temperature and precipitation: Deciduous broadleaved forest Evergreen needleleaved forest Portland, Oregon Erie, Pennsylvania 180 120 100 80 20 10 0 0 0 20 40 20 0 Precipitation Temperature 10 30 60 160 140 120 100 80 60 30 40 20 J F M A M J J A S O N D J F M A M J J A S O N D Month Month Temperature (C) 140 Precipitation Temperature Temperature (C) 160 Precipitation (mm) Precipitation (mm) 180 (1) By explicitly incorporating the seasonal interactions of energy and water, the climatic water balance is biologically more meaningful than traditional climatic metrics. 150 mm Water Potential evapotranspiration Water supply (rain plus snow melt water) 100 50 0 1 J F2 3 M 4 A 5 M J6 7J Month 8 A 9 S 10 O 11 N 12 D Actual evapotranspiration (AET) reflects the simultaneous availability of biologically usable energy and water, and is therefore an index of site potential for productivity. 150 mm Water Potential evapotranspiration Water supply (rain plus snow melt water) 100 S D 50 AET 0 1 J F2 3 M 4 A 5 M J6 7J Month 8 A 9 S 10 O 11 N 12 D Climatic water deficit (D) is evaporative demand that is not met by available water, and is thus an index of potential effects of drought stress on plants. (Don’t confuse with soil water deficit!) 150 mm Water Potential evapotranspiration Water supply (rain plus snow melt water) 100 S D 50 AET 0 1 J F2 3 M 4 A 5 M J6 7J Month 8 A 9 S 10 O 11 N 12 D Surplus is an index of biologically unusable water – excess water that leaves a site, through runoff or subsurface flow, without being evaporated or transpired. mm Water 150 100 S D 50 AET 0 1 J F2 3 M 4 A 5 M J6 7J Month 8 A 9 S 10 O 11 N 12 D (2) The climatic water balance (AET and D) is better correlated with the distribution of vegetation types. Mean annual precipitation (mm) Example: North American coniferous and deciduous forest distributions. 2500 2000 1500 1000 500 Deciduous forest Coniferous forest 0 0 5 10 15 20 Mean annual temperature (C) (2) The climatic water balance (AET and D) is better correlated with the distribution of vegetation types. 2500 2000 1500 1000 500 Deciduous forest Coniferous forest 0 0 5 10 15 20 Mean annual temperature (C) Annual actual evapotranspiration (mm) Mean annual precipitation (mm) Example: North American coniferous and deciduous forest distributions. 1400 1200 1000 800 600 400 200 Deciduous forest Coniferous forest 0 0 100 200 300 400 Annual climatic water deficit (mm) Stephenson 1998, J. Biogeogr. (3) The climatic water balance exhibits these properties across more than ten orders of magnitude of spatial scale. North America Stephenson 1990, Am. Nat. Stephenson 1998, J. Biogeogr. (3) The climatic water balance exhibits these properties across more than ten orders of magnitude of spatial scale. Sequoia Nat’l Park Stephenson 1998, J. Biogeogr. Summary -- the superiority of the climatic water balance in ecological studies: • More biologically meaningful. • Better correlated with biological processes and vegetation and species distributions. • Exhibits these properties across >10 orders of magnitude of spatial scale. Thesis: Consideration of the climatic water balance improves our ability to understand and forecast the effects of climatic changes on the biological world. (1) The superiority of the climatic water balance in ecological studies. (2) In a changing climate, increasing water availability will NOT counteract the effects of increasing evaporative demand. … And why soil moisture may be less important than you think! (3) Don’t forget AET! Mean annual temperature (C) At continental and global scales, most authors have treated metrics of energy and water supplies as orthogonal axes. That is, they recognize that water does NOT counteract the effects of energy. Mean annual precipitation (cm) Whittaker 1975 Elevation Yet at local to regional scales, some of those same authors additively combine metrics of energy and water supplies into a single “moisture scalar,” thus assuming that the effects of water and energy counteract one another. E.g., Whittaker’s moisture scalar combines metrics related to evaporative demand (slope aspect and exposure) with metrics related to water availability (slope position and proximity to water). “Topographic moisture gradient” Whittaker & Niering 1965, Ecology For understanding and forecasting plant distributions, we care less about soil moisture than the dynamics that gave us that soil moisture. “Normal” conditions Plant response Wet Dynamics of evap. demand and water supply Dry Water level For understanding and forecasting plant distributions, we care less about soil moisture than the dynamics that gave us that soil moisture. Drought (low precipitation) Wet Dry For understanding and forecasting plant distributions, we care less about soil moisture than the dynamics that gave us that soil moisture. Add water Wet Dry For understanding and forecasting plant distributions, we care less about soil moisture than the dynamics that gave us that soil moisture. Normal precip, but high evaporative demand Wet Dry For understanding and forecasting plant distributions, we care less about soil moisture than the dynamics that gave us that soil moisture. Add water Wet Dry Consider these scenarios in terms of annual AET and D. The colored dots represent species’ climatic niches; dotted diagonal are isolines of constant PET. Annual AET (mm) 200 100 0 0 100 200 Annual Deficit (mm) 300 Because PET = AET + D, drought (low precipitation) can only result in increases in D that are precisely offset by identical declines in AET. Annual AET (mm) 200 100 0 0 100 200 Annual Deficit (mm) 300 Adding the right amount of water can put the site back to exactly where it started in climate space. Annual AET (mm) 200 100 0 0 100 200 Annual Deficit (mm) 300 In contrast, because PET = AET + D, increasing PET must increase AET, D, or both. That is, the effects of changing PET are nearly orthogonal those of changing water availability … Annual AET (mm) 200 100 0 0 100 200 Annual Deficit (mm) 300 … so that adding water CANNOT put the site back to exactly where it started in climate space ... Annual AET (mm) 200 100 0 0 100 200 Annual Deficit (mm) 300 … and big enough increases in PET and water availability will move the site to another species’ climatic niche, even though soil moisture hasn’t changed. Annual AET (mm) 200 100 0 0 100 200 Annual Deficit (mm) 300 What are some implications for the effects of climatic changes on mountain ecosystems? Example: Unlike temperature changes, precipitation changes should not be expected to cause coordinated directional shifts in species’ elevations. PET declines sharply with increasing elevation. Evaporative rate (cc/day) 80 Santa Catalina Mtns, AZ, summer of 1911 60 40 20 South-facing slopes North-facing slopes 0 1000 1500 2000 2500 Elevation (m) Data from Shreve 1915 A spherical porous cup atmometer Livingston 1935, Ecology PET declines sharply with increasing elevation. Lo w at io n El ev h 100 H ig Annual AET (mm) 200 0 0 100 200 Annual Deficit (mm) 300 Thus, temperature-induced increases in PET will alter a site’s water balance to support species currently found at lower elevations. This is why we expect temperature-induced upslope migrations. Annual AET (mm) 200 100 0 0 100 200 Annual Deficit (mm) 300 At a given elevation, local water availability depends on things like soil depth, proximity to water, etc. (e Hi .g g so . de h ils ep ) Annual AET (mm) 200 Lo av ca ai l w la at bi e l it r y (e L .g o . w 100 so sha ils llo ) w 0 0 100 200 Annual Deficit (mm) 300 Declining precipitation will alter a site’s water balance to support species currently found at sites with lower water availability but at the same elevation … Annual AET (mm) 200 100 0 0 100 200 Annual Deficit (mm) 300 … and increasing precipitation will alter a site’s water balance to support species currently found at sites with higher water availability but at the same elevation. This is why we DON’T expect precip-induced elevational changes. Annual AET (mm) 200 100 0 0 100 200 Annual Deficit (mm) 300 These expectations are empirically supported by observations of species’ distributions along natural climatic gradients. Stephenson 1998, J. Biogeogr., Fites-Kaufman et al. 2007, TVC Thesis: Consideration of the climatic water balance improves our ability to understand and forecast the effects of climatic changes on the biological world. (1) The superiority of the climatic water balance in ecological studies. (2) In a changing climate, increasing water availability will NOT counteract the effects of increasing evaporative demand. … And why soil moisture may be less important than you think! (3) Don’t forget AET! Parting thoughts: “Moisture availability” is a slippery term of limited value, and can only be understood in two dimensions. E.g., does an increase in both AET and D yield a wetter or a drier environment? Arguably, both! In many (most?) cases it’s better to drop the concept of “moisture availability” in favor of explicit consideration of AET and D. Thank you for your attention!