Climate change is both an ecological and evolutionary event that - PowerPoint PPT Presentation

Climate change is both an ecological and evolutionary event that can force assisted migration and genetics-based ecosystem engineering Tom Whitham, Merriam-Powell Center for Environmental Research Not Bob Marley Heat map photo of & by Tom Whitham

SEGA site at The Arboretum at Flagstaff – A network of 10 common gardens along an elevation gradient to develop solutions to global change Part 1 - How locally adapted are plants? Importance - The more locally adapted, the greater the impacts of climate change.

( sega.nau.edu) $4.5 million NSF/NAU, GO, and NGO Participants: USFS, NPS, BLM, BOR, TNC, AZ Game & Fish, Babbitt Ranches, Grand Canyon Trust, & The Arboretum at Flagstaff Illustration by Paul Heinrich If plants must move to survive future climate conditions, how do we decide on which ecotypes and genotypes to use in restoration projects? SEGA network provides next generation genetics-based infrastructure to scientifically make such decisions.

Geographical Distribution of Ecotypes Geographical Central California adapted ecotypes Ecotype have evolved in Utah High Plateau response to Ecotype environmental differences across the range of P. fremontii . Sonoran Desert Ecotype Structure Analysis based upon molecular marker Ikeda et al. 2017 Global Change Biology

Utah High Plateau Ecotype Central California Ecotype Sonoran Desert Ecotype Most Predictive Environmental Variables Bio.15 – Precipitation seasonality Bio.6 - Max temp of warmest month Bio.11 – Mean temp coldest quarter Using genetically informed ecological niche modeling (gENM) with Maxent, we found that the regions occupied by different ecotypes will shift with projected climate change and will diverge spatially even more than their current distributions (Ikeda et al. 2017 Global Change Biology).

Genetics-based models are up to 12x better at predicting ecoregion test points Genetics No Genetics Genetics No Genetics 0 -2 Predictive accuracy -4 Binomial Probability (ln) -6 -8 -10 -12 -14 -16 Utah Central Sonoran -18 High Plateau California Valley Desert California Central Valley Sonoran Desert Utah High Plateau Ecoregion Ecoregion Ikeda et al. 2017 Global Change Biology

Reciprocal common gardens show finer scale local adaptation within the Sonoran desert ecotype Reciprocal TNC Dugout Ranch, Canyonlands Common Cooler Garden – MAT 10.7 ° C Gardens AZG&F Horseshoe Ranch Intermediate – MAT 17.2 ° C BLM Mittry Lake, Yuma Hot Garden – MAT 22.8 ° C Location map of 16 provenance collection sites (leaf icon) of Populus fremontii and the three common garden locations (leaf with circle). The central garden is also a collection site. The shading corresponds to the degree-days above 5 ° C (DD5) throughout the region: red represents high DD5, blue low DD5. (Cooper et al. unpub.)

March 8, 2017 – Photos by Tom Whitham Agua Fria Creek Populus fremontii field trial Indian Creek 4600 tree common garden on Arizona Game & Fish Dept. lands at Horseshoe Ranch surrounded by Agua Fria National Monument.

Hot Garden Intermediate Cool Garden Long Growing Season Short Growing Season Populations are locally adapted within an ecoregion Population level mean (+/- 1 SE) survival correlations with bud set date in each of the three common gardens. Populations are colored by the mean annual temperature (MAT ° C) of their source provenance. In Yuma, survival is highest in the hotter source populations and is positively correlated with later bud set. The opposite is true in the coldest Canyonlands garden. From Cooper et al. 2018 Global Change Biology (in press).

Fremont cottonwood P. fremontii Geographic mosaic of divergent functional trait selection Divergent selection in quantitative traits affected by climate Qst-Fst of plant functional traits (bud phenology, height, DRC, SLA) across the three common gardens (Cooper et al. unpub. data). Same has been shown for narrowleaf cottonwood, P. angustifolia (Evans et al. 2016)

Part II. Genetic differences in the functional traits of plants affect community structure and ecosystem processes. Importance – If climate selection is non-random, then community structure and ecosystem processes will change.

Genetic “footprints” of trees can be large: The genetic links between terrestrial and aquatic ecosystems. Intraspecific differences in cottonwoods affect stream macro-arthropod communities. Populus angustifolia Emergence trap photo by Zacchaeus Compson

1.0 996 0.8 1008 1012 0.6 1017 1020 0.4 RM2 T15 0.2 Axis 2 0.0 -0.2 -0.4 -0.6 -0.8 -1.5 -1.0 -0.5 0.0 0.5 1.0 1.5 Axis 1 Different genotypes of P. angustifolia support different stream macro-invertebrates. Compson 2016 Ecosphere

Community Phenotypes Frequency Traditional Phenotype of Resistance Aphid survival AFLP genetic markers Ecosystem Phenotypes Whitham 1989 Science, Dickson & Whitham 1996 Oecologia, Schweitzer et al. 2006 Oikos, Keith et al. 2010 Ecology, Zinkgraf et al. 2016 J. Insect Physiology

Genetic Linkage Map of Populus lg1 lg1 lg6 lg6 QTL Mapping Insect Resistance Leaf Chemistry Architecture Phenology lg8 lg8 lg8 Salicortin lg18 Spring Leaf lg12 Flush Resistance to P. betae Second Year Branch Number Condensed Tannin Woolbright et al. 2008 Heredity Matt Zinkgraf et al. 2016 J Insect Physiology Woolbright et al. 2018 Ecology & Evolution

Illustration by Victor Leshyk Why do different genotypes support different communities? Many functional traits result in individuals being very different from one another and affecting other traits such as diversity, stability, and network structure.

San Francisco Peaks & Wupatki National Monument view from Navajo Nation Photo by Tom Whitham Communities are far more linked than previously envisioned.

Lichen 2010 P ! 0.05 Lichen 2010 Lamit et al. 2015 A 0.05< P <0.1 P ! 0.1 Journal of Ecology Fungal Leaf Ectomycorrhizal Leaf Pathogens 2010 Phyllosphere EM Fungi 2006 Pathogens 2010 Trunk Fungi 2006 Soil Fungal Leaf Soil Bacteria Leaf Pathogens 2009 Soil Bacteria 2004 Pathogens 2009 2004 Leaf Modifying Leaf Modifiers 2010 Soil Fungi Soil Fungi 2004 Arthropods 2010 2004 Populus angustifolia Twig Endophyes Endophytes Endophytes 2006 2006 The network of correlated communities is defined by individual tree Lichen 2010 Lichen 2010 B genotypes – the importance of maintaining network structure in restoration. Ectomycorrhizal Fungal Leaf Leaf Pathogens 2010 EM Fungi 2006 Fungi 2006 Pathogens 2010 Community-genetic correlations - changes in the composition of one community among plant genotypes that are mirrored by changes in the composition of another community.

Fremont cottonwood on the Little Colorado River - Photo by Tom Whitham Part III. Climate change creates a mis-match of once locally adapted plants and the new environment that affects plant survival, biodiversity, and mycorrhizal mutualisms.

Climate change results in mis-matches with the local environment. In just 25 years plant hardiness zones have shifted northward and upward in elevation by one zone.

Photos by Hillary Cooper (top) & Tom Whitham (bottom) Cottonwood mortality on Bill Williams River National Wildlife Refuge – March 28, 2017 Stand die-off due to record drought – plants are no longer adapted to the local environment. As the local stock dies out, what populations and genotypes should be used in restoration?

Record drought of 2002 – An evolutionary event that changed the genetic structure of the tree population. Sthultz et al. 2009 Photo by Tom Whitham Sept 15, 2004 - North of Global Change Biology San Francisco Peaks, AZ

Diverse arthropod community on pinyons negatively affected by drought

40 155 arthropod species Total richness per tree 30 Within a site, increased 20 stress negatively affects arthropod diversity. 10 Species Richness 0 Stress Index = (standardized r 2 =0.53, p<0.0001 branch dieback + number of needle cohorts + 0.0 0.5 1.0 1.5 2.0 2.5 Stress Index radial trunk growth) 25 Total abundance per tree 20 15 10 5 Abundance 0 r 2 =0.53, p<0.0001 Stone et al. 2010 Oecologia -5 0.0 0.5 1.0 1.5 2.0 2.5 Stress Index

Part IV - Solutions to the mis-match EVOLVE, ACQUIRE BETTER MUTUALISTS, MOVE or DIE Painted Desert from SP Crater Photo by Tom Whitham

Genetics-based responses to climate change Genetic Variation in Plasticity Plasticity Genetic A two-environment reaction norm showing the components of phenotypic variation of four genotypes: G = trait variation due to population genetics within a single environment, E = trait variation due to change in environment (plasticity), GxE = the variation in plasticity among genotypes. Phenotypic variation (V P ) = V G + V E + V GxE . From Cooper et al. 2018 Global Change Biology (in press).

Rapid Evolution with Drought Drought susceptible trees were >3X more likely to die during the 2002 record drought than drought resistant trees. 80 70 60 % mortality 50 40 30 20 10 0 Drought Drought Susceptible Resistant Pinyons Pinyons Gehring et al. 2014 Molecular Ecology