Community and Evolutionary Consequences of Record Drought in the Southwest Gery Allan – molecular systematics Randy Bangert – biogeography Samatha Chapman – nutrient cycling Neil Cobb – insect ecology Steve DiFazio – molecular ecology Dylan Fischer – ecophysiology Robert Foottit – molecular systematics Alicyn Gitlin – restoration ecology Steve Hart – ecosystem/soil ecology Paul Heinrich – public outreach Paul Keim – microbial/bacteria genetics Karla Kennedy – resotration ecology Zsuzsi Kovacs – mycorrhizal ecology Rick Lindroth – chemical ecology Jane Marks – aquatic ecology Nashelly Meneses – ecological genetics Brad Potts – population genetics Jen Schweitzer – ecosystems Steve Shuster – theoretical genetics Chris Stulz – population ecology Randy Swaty – conservation biology Amy Whipple – ecological genetics Gina Wimp – community ecology Scott Woolbright - molecular genetics Joe Bailey – community ecology Brad Blake – restoration biology Aimee Classen – nutrient cycling Ron Deckert – endophyte ecology Eck Doerry – bio-informatics Kevin Floate – insect ecology Catherine Gehring – microbial ecology Allen Haden – aquatic ecology Kristin Haskins – mycorrhizal ecology Barbara Honchak – ecological genetics Art Keith – insect community ecology George Koch – ecophysiology Carri LeRoy – aquatic ecology Eric Lonsdorf – genetic modeling Greg Martinsen – ecological genetics Becky Mueller – plant ecology Brian Rehill – chemical ecology Crescent Scudder – plant demography Adrian Stone – insect communities Richard Turek – statistics Talbot Trotter – dendrochronology Tom Whitham – ecology Stuart Wooley – phytochemistry Tong-Ming Yin – quantitative genetics Key Issues 1. Climate change alters community structure and negatively affects biodiversity. 2. Drought impacts on dominant and keystone species are most important because they are community drivers. 3. Some vegetation types are far more likely to be affected than others. 4. Drought is likely to be an evolutionary event. 5. There are likely to be major surprises with drought events that cannot be predicted. 2200 years of climate history argues major switches in selection pressures from wet to dry periods that could affect biodiversity and rapid evolution. Grissino-Meyer 1996 Expansion of junipers (Juniperus monosperma) into grasslands 1899 1977 Enchanted Mesa, New Mexico Allen et al. 1998 Pinyon mortality North side of the San Francisco Peaks, AZ Photo mosaic by Paul Heinrich taken December 2003 Photo by Tom Whitham Sept 15, 2004 - North of San Francisco Peaks, AZ Photo by Tom Whitham Sept 15, 2004 - North of San Francisco Peaks, AZ 2000 75 UT km2 2003 CO 2001 332 km2 2002 1,377 km2 km 2003 10,406 km2 Total 12,191 km2 AZ NM Pinyon + Ponderosa Pine Mortality - Compiled by the National Forest Health Monitoring Program Slope Function Aspect Function Elevation Function = Low Risk = Moderate Risk = High Risk = Known High Pinyon Mortality Sites Model correctly intersects known high pinyon mortality sites with 75% accuracy. ~10% of error is from low resolution DEM ~15% of error is from undocumented high pinyon mortality sites + unknown error. Trotter et al. 2005 Lightning strikes twice in the same spot. Fig. 1 Stand-level % Pinyon Mortality in 2002 80 r2 = 0.504 F = 9.15 p = 0.01 60 40 20 0 0 20 40 60 Stand-level % Pinyon Mortality in 1996 Mueller et al. 2005 80 Larger pinyons are far more likely to die than smaller ones. Fig. 2 120 100 r2 = 0.712 F = 69.4 p < 0.001 % Mortality 80 60 40 20 0 0 10 20 Size Class (1.0 cm) Mueller et al. 2005 30 Mean Arthropod Richness / Tree On Trees Growing Under Low Stress 14 B High Stress Low Stress B 12 10 8 A A B 6 4 A 2 Mean Arthropod Abundance / Tree 0 1B: Overall Mean Arthopod Abundance Is Highest Figure Predators Parasites OnHerbivores Trees Growing Under Low Stress High Stress Low Stress 600 B 40 20 A In an arthropod community of 266 species, species richness is 2 X higher and abundance is 12 X higher in low stress sites than high stress sites. B A A B Trotter et al. 2005 0 Herbivores Predators Parasites Figure 2: Canopy Arthropod Community Composition Differs Significantly Between Trees Growing InCommunities High and Low Stress Changes Arthropod Stress Environments 0.6 High Stress Sites 0.4 Y Axis 0.2 High Stress 1 High Stress 2 High Stress 3 Low Stress 1 Low Stress 2 Low Stress 3 0.0 -0.2 -0.4 -0.6 -0.6 Low Stress Sites -0.4 -0.2 0.0 X Axis 0.2 0.4 0.6 Trotter et al. 2005 Watering experiment conducted by Crescent Scudder to test for community release. Photo taken May 23, 2003 Mean Athropod Abundance on Arthropod Non-watered vs Abundance Watered Pinyons 40 p=.028 30 Watered Pinyons Non-watered Pinyons 20 10 0 Watered Control Mean Arthropod Richness on Non-watered vs Watered Pinyons 20 Species Richness p=.031 Supplemental watering increases arthropod abundance and species richness. 15 10 Scudder unpub. data 5 0 Watered Control Watered and control trees support different arthropod communities (Scudder et al. 2005). 2003 Community Composition of Watered and Control Trees Axis 2 Watered Trees Control Trees Watered Control Axis 1 Ectomycorrhizae on fine root of pinyon pine Environmental Stress Shifts Ectomycorrhizal Community O'Neill Crater RFLPs R = 0.652 P < 0.00001 Low Mortality Sites High Mortality Sites Swaty et al. 2004 Ecology Low Mortal High Morta Tree Rings Predict Arthropod Species Richness 30% Richness vs. Ave 97-01 40 Species Richness per Tree Y Data 30 20 10 0 0 500 1000 1500 2000 2500 X Data Average Tree Ring Width 1997-2001 Ave 97-01 vs 30% Richness Plot 1 Regr Adrian Stone unpub. data 3000 Moth Resistant Pinyon These phenotypes are genetically based and have extended phenotypes that have community consequences. Susceptible Pinyon Photo by Tom Whitham Switch in performance - Moth resistant trees 3X more likely to die during 2002 drought than moth susceptible trees. 80 % mortality 70 60 50 40 30 20 10 0 Stulz et al. unpub. data Moth Resistant Pinyons Moth Susceptible Pinyons Scales associated with reversals • • • • Number of species Time Spatial scales Any mix of above Interactions Increase With the Addition of Factors % of Significant Interaction Terms 100 Observed Total Observed Significant Interactions Observed ObservedReversals Reversals 80 60 40 20 0 0 1 2 3 4 Number of Statistical Factors (Species, Time, Space) # of Statistical Factors Bailey & Whitham 2005 The extended phenotypes of moth resistant and susceptible trees affect a diverse community of about 1000 species. Whitham et al. 2003 Ecology Drought impacts most dominant plants and their dependent communities. Photos by Tom Whitham & Alicyn Gitlin Trees like cottonwoods should be of special concern due to low coverage. Gitlin et al. unpub. data Landscape coverage (%) Population mortality (%) Mortality of dominant plants at 20 randomly selected sites for each species within a 80km radius of Flagstaff. 60 50 P < 0.0001 a 40 30 b b 20 10 b b c b 60 0 50 40 30 20 10 0 (0.16) Juni. Manz. Asp. Pond. Cotton. Piny. Dominant plant species Summary 1. Through its effects on community drivers (e.g., dominant trees and keystone herbivores), drought alters community structure and negatively affects biodiversity. 2. Some dominant vegetation types are far more likely to be affected than others. 3. Switches or unexpected outcomes are likely (e.g., insect resistant trees are more likely to die, which in turn results in a major community shift). 4. Extreme drought is a bottleneck event that is also likely to be an evolutionary event. Management & Research Issues 1. In the absence of long-term community-level studies, fundamental errors in interpretation are likely due to the high probability of switches. 2. Need to minimize human impacts that exacerbate the effects of drought (e.g., water diversions that make a 100 year drought into a millennium-level drought). 3. Need special emphasis on rare habitat types that are especially sensitive to drought (e.g., riparian habitat and springs). 4. Marginal or edge habitats that suffer chronic stress can be barometers of change and may be crucial to preserve as sources of extreme genotypes that may be best adapted to changing environments. 5. Restoration with local genotypes may not best strategy if climate change continues (e.g., drought adapted genotypes from lower elevations should be included in a diverse genetic seeding program). 6. Need to better interface climate change models with community change models. DIREnet (Drought Impacts on Regional Ecosystems Network) coordination and synthesis of ecological research on drought effects and the potential role of global climate change. http://www.mpcer.nau.edu/direnet/